\r\n\tRadiation monitoring deals with the sampling and measurement of different products found in different radiation pathways from the environment ending with consumption in humans. Gamma-spectroscopy is the main tool for measurement of these radiations.
\r\n
\r\n\tThe aim of this book is to investigate the radionuclide concentrations in the most consumable food products, air, water and soil. Particularly, it is essential to investigate the radiations level in the surroundings of a nuclear facility.
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1. Introduction
The life of ants as ectothermic organisms is closely connected with the seasonal variations of ecological factors, such as temperature, rainfall, humidity, availability of food, etc., occurring during the year. Certain climatic changes usually manifest even in tropics and cause the corresponding modifications in behavior and development of ants and other insects, but they are much more pronounced on the north. In the regions with temperate and boreal climate, the annual changes of climatic conditions possess a hard impact to the life cycles of all organisms living there. In different ant species, various annual cycles of behavior arise. They have been described in many reviews (for example, see [1, 2]). However, the seasonality of the behavioral cycle in ant colonies is strictly subordinated to the seasonality in developmental processes such as oviposition or larval rearing. Each ant species demonstrates certain annual developmental cycle because the processes of the colony development cannot proceed without interruption during the whole year.
To optimize the survival and growth of ant colonies, the entire warm season should be used for larvae rearing and for production of the greatest quantity of new adults. That is why the brood development should start in spring as earlier as feasible and prolong as longer as possible in late summer. At the same time, only the brood stages that are able to survive during the winter should occur in ant colony by the end of autumn. To ensure this, the special physiological mechanisms evolved which provide synchronization of the colony development with the yearly climatic periodicity.
The first investigations devoted to seasonality in the development of ants were performed by Yozhikov who studied the phenology of the development in several ant species from Central Russia [3]; by Headly who characterized the seasonal development of two Leptothorax species and Aphaenogaster fulva aquia in North America [4, 5]; by Talbot who carried out the phenological studies on two North American Myrmica species [6]; and by Eidmann who described the hibernation in ants and mentioned that the Formica species overwinter without a brood [7].
Despite extensive studies of arthropod dormancy and seasonality, myrmecologists paid very little attention to the role of seasonality in ant ecology, as well as in the evolution of the life cycles of these social insects. Literature on this topic is not rich. Quite a few publications specifically devoted to seasonal development and the phenology of ants. As a rule, such data can be found in the investigations dealing with the biology and ecology of certain species. Even less frequently, the subject of the study was the regulation of annual cycles of the ant development. Most often, these problems were affected accidentally and stood in the background and pushed back by the main aims of the study (for example, see [8]). The seasonal development of the ants Myrmica was studied to the greatest extent, mainly due to the works of M. Brian (for reviews, see [1, 8, 9, 10]).
This paper contains a review of literary and proprietary data on the structure, diversity, adaptive features and evolution of seasonal cycles and strategies of the seasonal development in ants. We have studied the seasonal cycles in more than 80 species belonging to more than 20 genera from different regions of Russia and the former USSR, ranging from warm temperate to cold temperate and boreal climatic zones. Our field and laboratory studies have allowed us to map the diversity of annual cycles, to reveal the underlying ecophysiological and social mechanisms of control and to develop ideas on possible pathways in the evolution of the seasonal cycle in ants.
The main study methods we have used were laboratory experiments and field phenological observations. In experimental studies, the ant colonies were divided into fragments each consisted of workers, queens and the brood. In the case of a small number of workers in the colony or for monogynous species with a single queen in the nest, entire natural colonies were used. Colony fragments (or the whole colonies) were kept in artificial plastic nests, which were randomly distributed over various experimental regimes (different photoperiodic conditions and constant temperatures or thermal periods) and experimental regimes were maintained in special thermostats. Our methods of laboratory cultivation of ants provided the opportunity to observe and to study all stages of annual cycle including the overwintering in a refrigerator under the temperatures of 3–5°C.
2. Peculiarities of seasonal development in ants as social insects
In social insects, in addition to the life cycles of individual individuals, there is a life cycle of the colony, as an integrated, superorganismic system. It consists of the processes of the development of individuals, but it is not equivalent to a simple summation of them. Social regulation mechanisms arise in the evolution and are realized through interactions between members of the colony. They control the physiological state and the development of individuals, depending on the ecological situation and colony needs. Such “collective regulation” of development is absent in solitary species and largely determines the specificity of seasonal development in ants [11, 12, 13, 14, 15, 16].
Colonies of ants are not only perennial but usually have unlimited life cycle under favorable conditions of environment [17]. Not only queens but even workers can survive for several years. In monogynous species, all workers and the brood are the offspring of the only queen. So while the queen is alive, all the population of a colony can relate to one and the same genetic generation during consistent years. In polygynous species with several queens in a nest, all individuals inhabiting a single colony may pertain to various generations which overlay each other. However, the seasonal life cycle of the colony is not associated with differences arise between generations. It embraces the regular seasonal variations in physiological state of all individuals in a colony which entail the orderly changes in behavioral and developmental patterns. Therefore, we determined the seasonal life cycle of an ant colony as the annual cycle of physiology, behavior and development [18].
After the spring awakening, the queen commences laying eggs, the development of larvae begins and pupae appear. There are workers and reproductive individuals among new adults. Oviposition and brood development continue throughout the warm period of the year and cease in the autumn, when insects begin to prepare for wintering, go to special shelters and spend the winter in inactive state. The annual cycle of colony development is a collective and highly organized process and includes the individual development of immature stages (the brood) and regular seasonal changes in the physiological state and reproductive activity of adults (workers and queens).That is why the growth and development and the beginning and termination of diapause in ants can be considered and studied both in individual level and at the level of the entire colony. And this is not the same thing, since all these processes are under the control of the mechanisms of social regulation and integrated reactions of the colony to the changes in environmental conditions. In this connection, diapause of ant larvae can be (and usually happens) facultative at the level of an individual, but at the same time obligatory at a colony level (in endogenous-heterodynamic species).
3. Homodynamic type of seasonal development
On the general background of an insufficient study of the phenology in ants, the tropical regions of the Earth are especially prominent. Analysis of meager data shows that at any time of the year, in the nests of most tropical species studied, all the stages of ontogenesis from the egg to the pupa are present, and development retardations are absent. Such continuous all-year round development without the obligatory onset of periods of physiological dormancy we call homodynamic, using the terminology of E. Roubaud [18, 19, 20, 21].
However, homodynamic species often have a certain seasonal structure of the annual cycle: on a general background of continuous development, there may be significant seasonal fluctuations in the number of certain ontogenetic stages, as well as seasonal association of the rearing of alates and nuptial flights. Thus, in Cataulacus guineensis from Ghana, the brood population has two maxima—in May and in September, the alate females and males are numerous in the nests in July–October, and the rest of the time there are few or none alates at all [22]. The larvae of alate reproductives of Camponotus sericeus in India develop from October to July, and alate females and males can be found in nests all-year round, but their nuptial flight occurs in September–October [23, 24]. Colonies of the tropical ant Paltothyreus tarsatus (Republic of Côte d’Ivoire) grow alate females in spring, and males in autumn; reproductives live in maternal nests for a long time and leave for mating only in February [25]. In definite periods of the year, alates of Anoplolepis longipes are grown up in Papua New Guinea [26] and in the Seychelles [27] and alates of Camponotus detritus in Namibia [28]. The factors that cause such seasonality in tropical species are not yet clear enough.
In addition, periodicity of oviposition was noted for a number of species. For example, in the natural colonies of Anoplolepis longipes, Cataulacus guineensis and species of the genus Rhytidoponera [29], there are two egg abundance maxima throughout the year. Rhythm of oviposition was also found in a tropical species Linepithema humile, widespread in Southern France, both in natural and laboratory conditions [30]. The periodicity of egg laying is also a characteristic feature of the nomadic ants of the tropical subfamily Dorylinae and especially of the Neotropical species. In all species, the cyclic brood rearing is clearly associated with behavioral cycles: oviposition occurs during stationary phase of reproductive cycle that alternates with nomadic phase, during which the ants feed the larvae that emerge from the eggs [31, 32, 33].
The homodynamic development of some tropical ants was observed in the laboratory. G. Terron maintained colonies of the African species Tetraponera anthracina at 25–26°C for several years and reported that development of the brood occurred continuously and reproductives periodically appeared [34]. But he observed alternating periods of egg laying and reproductive dormancy of queens. The ant Monomorium pharaonis, a widespread synanthropic species, which was imported from tropical Africa to Europe and North America, now inhabits many heated buildings. It was found that in the nests of this species under optimum conditions (27–30°C, relative to 60–65%), brood developed continuously with cyclic rearing of alate reproductives at regular intervals [35, 36, 37, 38, 39, 40, 41]. We also kept several colonies of M. pharaonis found in St. Petersburg in the laboratory for 2 years and revealed that at any temperatures above the threshold, which is 17.7–17.8°C in this species [42], brood development occurred without any delays. However, at a near-threshold temperature of 17°C, the mortality rate of all brood categories, as well as of adult ants, sharply increased, and the colonies died within a month. Thus, this tropical species cannot tolerate even short periods of temperature decrease, which confirmed by the available data on its habitation in Europe only in heated buildings [43].
We observed homodynamic development in two species from tropics: Pheidole sexspinosa from the Tonga archipelago and the Tetramorium semillimum from the Seychelles. We kept them in the laboratory for more than a year, varying the temperature and photoperiod within acceptable values, i.e. within those that can be observed in the natural habitat of these species (18–25°C, 10–16 hours of light per day). All this time, there was a continuous development in the colonies.
It is well known that typically tropical insects cannot survive for a long time at temperatures well below the optimum and, especially, below the developmental threshold [44]. Therefore, it can be argued that the tropical ants in their majority are not adapted to survive during the cold periods of the year. However, some ant species can demonstrate continuous all-year round development even in subtropical environments. For example, in the central regions of Texas (USA), Pseudomyrmex sp. occupies the stems of a mimosa, and all ontogenetic stages are present year round, but in different amounts: eggs and larvae are numerous in winter and pupae in spring and summer [45]. Thus, we can assume that the development and pupation of larvae of this species continues in winter, but much more slowly than in summer. This decrease of brood developmental rate leads to a significant decline in the number of pupae in wintering nests. According to Rissing, new workers of Messor (Veromessor) pergandei in Arizona (USA) appear from pupae also throughout the year [46].
4. Heterodynamic cycles of development
Annual developmental cycles of ant colonies, in which a diapause arises naturally, we call after Roubaud heterodynamic [18, 19, 20, 21]. They have a distinct seasonal structure: the period of diapause (the phase of dormancy) is regularly replaced by the period of development (the active phase of the cycle), after which a new period of diapause occurs, and so on. Generally, the phase of dormancy in the annual cycle coincides with the period of unfavorable climatic and (or) food conditions. It is characterized by a lack of larval development and, as a rule, of queen oviposition and the presence in the nests of only certain (usually diapausing) brood categories. During the active phase of the annual cycle, eggs are laid and the larvae are reared. Ants grow up new workers, as well as alate females and males.
4.1. Quasi-heterodynamic development
Some ant species recently penetrated from tropics or subtropics to areas with a warm temperate climate and successfully settled there. Two species of fire ants, Solenopsis richteri and S. invicta, were imported to the United States from South America: the first one, around 1920, and the second one, in the early 1940s [47, 48]. S. invicta significantly expanded its original range fairly far north and is now prevalent in most southeastern states [48, 49]. The distribution of S. richteri is limited mainly to the northern parts of Alabama and Mississippi and the southern part of Tennessee [50].
Several authors have shown that in southern United States (Mississippi, Florida, South Carolina, etc.) both fire ant species remain essentially homodynamic: eggs, larvae and pupae of workers are present in their nests all-year round [48, 51, 52]. However, the number of immature stages varies considerably during the year, and in winter, it is very small. In February–March, the number of eggs increases sharply and new larvae develop and begin to pupate in April (workers) and May (alates). At this time, the quantity of brood categories is maximal and reaches 40–45% of the entire biomass of the colony. Reproductive individuals appear from pupae in June and fly out of the nests at least five times during the summer. The second small peak of the brood population (up to 35% of the biomass) in the nests coincides with the first cooling in September–October. At this time, all larvae and pupae belong only to the caste of workers. In November–December, the number of larvae and pupae sharply decreases and reaches a minimum (less than 2% of the biomass of the colony) in January. Thus, the number of immature stages in fire ants directly depends on the environmental temperature.
It has been determined that in the most northern populations of S. invicta, oviposition and brood development are suspended in the coldest months, i.е. eggs and pupae are absent in the nests in winter [48, 53]. It can be assumed that there is no diapause of larvae in fire ants and larval development ceases only when the ambient temperature falls below the developmental threshold, which is about 17°C according to laboratory experiments [54]. We can also assume that many eggs, larvae and pupae (as well as adult ants) die during the winter in northern populations of this species. The winter mortality of S. invicta workers was noted in several works (for example, [55]). Nevertheless, the colonies successfully survive in these conditions. Consequently, these ants already have some physiological adaptations that allow them to tolerate a sufficiently long stay at low positive temperatures.
Such tropical species, adapted to live in regions with cold winters without forming real diapause, we call quasi-heterodynamic [18]. They are characterized by the potential for unlimitedly long development under favorable conditions inherent in homodynamic species. The development of their brood ceases only at temperature below the developmental threshold and ants spend the winter in a quiescent (cold coma) state suffering from more or less strong mortality, but in general the colonies overwinter successfully.
The Argentine ant Linepithema humile demonstrates another example of quasi-heterodynamic development. This tropical species was imported from South America and now widespread in many parts of the world. Its phenology, investigated in the USA [56] and in Europe [30], is very similar to the seasonal development of fire ants. In the southern part of California, only a few larvae of workers and very few eggs can be found in overwintering nests of this species. At this time of year, more than 90% of the colony biomass is made up by adult ants. The new seasonal cycle begins in late February–early March, when the queen starts to lay eggs. The larvae hibernated in the nests complete their development by the middle of March. In summer, brood makes up about 50% of the colony biomass, but in October its number decreases sharply and gradually reaches its minimum by December. On the southern coast of France, the Argentine ant has similar phenology. Most small larvae hibernate in the nests, but in small numbers, and there are also some medium and large larvae and very rarely eggs. Renewal of development is observed in March.
Since the fire ants and Argentine ants recently permeated to the areas with cold winters, we tend to think that they do not yet have diapause. However, it has been experimentally determined that only the overwintered colonies of the Argentine ant can grow up a lot of alate females, which appear in the nests in the south of France at the beginning of the summer season [57]. This suggests that there are seasonal changes in the physiological state of colonies that are similar to diapause.
According to the literature date, many subtropical ants do not have any brood in their nests during the winter, for example, Ponera pennsylvanica in the state Missouri [58], Pogonomyrmex rugosus and P. subnitidus in Southern California [59] and Prenolepis imparis in the states Missouri, Ohio and Florida [60, 61, 62]. However, without experimental laboratory studies, it is impossible to conclude whether all of them are quasi-heterodynamic and overwinter without brood due to its death, or, conversely, they have a stable winter diapause of the queens, i.e. belong to the group of true heterodynamic species.
We discovered and investigated quasi-heterodynamic developmental cycles in several ant species living in regions with subtropical and warm temperate climates. In the colonies of Tetramorium nipponense and Pachycondyla chinensis, which were collected in the south of Japan and were kept in the laboratory at temperature of 25–27°C and photoperiod of 16 h of light per day, the oviposition of queens and the development and pupation of larvae did not cease during the year. After a gradual decrease in temperature, the colony successfully overwintered at 7–8°C. At the same time, however, all brood died, which probably occurs during the overwintering in natural conditions of the south of Japan.
In the experiments on Monomorium kusnezovi from Turkmenistan, the growth, development and pupation of larvae continued uninterruptedly at temperatures exceeding the developmental threshold of 20°C for eggs and 21.5°C for larvae and prepupae [42]. Thus, larvae of this species do not have diapause, which is a sign of quasi-heterodynamic seasonal cycle. At 20°C, the larvae ceased growth, but the oviposition did not stop, and by the beginning of overwintering, eggs and larvae of all ages were still present in the nests. However, eggs, as a rule, did not survive during hibernation in the laboratory, but the larvae hibernated more successfully. We assume that, in natural conditions, these ants go on hibernation with eggs and larvae, but eggs and part of the larvae die during the winter. In the middle of April in the Central Kopet Dag when we dug out the nests of M. kusnezovi, we always found small packets with eggs and larvae of younger instars and a small number of the third instar larvae lying separately, which already began to pupate at this time of a year. Obviously, these larvae overwintered in the nests but the eggs could appear in the early spring. Thus, it seems likely, though not completely proved, that a certain number of eggs can be preserved in the nests of M. kusnezovi until spring.
Two Pheidole species (P. pallidula and P. fervida) were collected in Turkmenistan and Russia, where they inhabit the regions with temperate climate. They also proved to be quasi-heterodynamic. P. pallidula is common in southern Europe, the Caucasus and Central Asia [63]; the distribution area of P. fervida includes Southeast Asia, Japan, southern Kurils and the south of Primorye [64]. In Primorye, P. fervida probably survived from the Tertiary period, when the climate there was much milder. In our experiments with both species, the queens continued to lay eggs and the larvae developed and pupated at any temperatures exceeding the developmental threshold (about 18°C for P. pallidula [42]). Any forms of diapause were absent. Under optimum conditions at 25–28°C, we observed continuous development for two or more years.
For P. pallidula from Turkmenistan, we found that when the temperature decreased to 20° C, the queen’s productivity declined significantly, but queens did not stop laying eggs, and the larvae developed and pupated. At a temperature of 17°C, which is only slightly below the developmental threshold for eggs and larvae, oviposition continued, but the eggs did not develop. Pupae and prepupae were the first to die, and then all brood stages gradually perished. Consequently, it could be assumed that in nature all brood in the nests of this species die even before the beginning of winter. Our field studies in Turkmenistan have shown that there was no brood in the nests of P. pallidula in early spring indeed. It appeared later from the eggs laid by queens. This is confirmed by the observations made by Passera [65, 66, 67] in the south of France.
P. fervida is somewhat better adapted to the temperate climate in the south of Primorye and its brood probably dies out only partially during overwintering. In our experiments with autumn colonies of this species, even at a temperature of 17°C and under short day conditions (10 hours of light per day), the queen continued oviposition, and larvae still pupated. We maintained one colony, in which there were originally more than a hundred larvae of different size, under these conditions during 7 months. Over this time, some of the larvae perished, but more than half of them pupated. This situation was strikingly different from the one we observed in similar experiments with truly heterodynamic ants: in most cases at 17°C, the development very quickly ceased due to the onset of diapause.
When we decreased the keeping temperature for the autumn colonies of P. fervida up to 10–12°C, the workers began to dismember and to throw out the prepupae and pupae from the nests and gradually destroyed them all. There were only eggs and larvae of all instars in the nests. However, part of the eggs and larvae of the first and second instar perished during the hibernation in the refrigerator at 3–5°C, while the older larvae overwintered more successfully. It would be extremely interesting to compare the population of the colonies of this species in late autumn and early spring to assess the ability of the larvae to survive during the winter in natural habitats. Probably, the similar regulation of the seasonal developmental cycle exists in Pheidole morrisi, which hibernates with larvae in the north of Florida [68].
4.2. True heterodynamic development
Most temperate and all boreal climate ants are true heterodynamic. They possess real winter diapause in their annual cycles (prospective dormancy) [18]. The presence of this diapause provides a more successful wintering by increasing the physiological resistance of larvae and adult ants to unfavorable winter conditions. In the literature, there is practically no data on the tolerance of developing and diapausing larvae, other developmental stages and adult ants to low temperature and other unfavorable environmental factors. Plateaux [69] noted that Temnothorax nylanderi in an active physiological state could not successfully overwinter: when he put summer colonies of this species into the refrigerator, eggs, pre-pupae and unpigmented pupae quickly died and began to rot, causing the death of the entire colony. Similar results were obtained in our experiments with colonies of Lasius niger, Lepisiota semenovi, Myrmica rubra, M. ruginodis and Plagiolepis compressus, which we put in the refrigerator at a temperature of about 5°C in summer. Eggs, pre-pupae and pupae died within 1–3 weeks, but larvae remained alive.
True heterodynamic seasonal cycles occur in the vast majority of ant species living not only in temperate and cold climates, but also in subtropics and even in tropics. The presence of long-term developmental delays was noted, for example, in all five species from the Rhytidoponera impressa group, widespread in the forests of Eastern Australia. These species demonstrated a strict seasonality of development: during the winter months, only small and medium-sized larvae and very rarely eggs were found in their nests. This situation was observed both in subtropical and tropical regions of Australia [29].
It is clear that for the occurrence of heterodynamic development in tropics, any seasonal changes in environmental conditions have to exist. Since the annual rhythm of the climate is usually quite distinct in the tropical regions, and its absence, on the contrary, is very rare situation, heterodynamic seasonal cycles should probably be widespread in tropical ants. The hibernation and diapause are widespread in tropical insects, but mechanisms of the regulation of heterodynamic cycles in tropics are far from understanding and explaining yet [70].
It should be assumed that heterodynamic development is more common for ants in subtropics, because the seasonal rhythm of the climate there is much more pronounced than in the tropical zone. Indeed, most of the subtropical species studied demonstrate the cessation of development in winter. Some of them do not have brood, while others overwinter with larvae: Camponotus kiusiuensis [71], C. nawai [72] and Leptanilla japonica [73] in southern Japan, Temnothorax monjanzei in Algiers [74] and T. melas on the Corsica island [75], Tapinoma minutum in New South Wales [76], Polyrhachis vicina in southern China [77, 78] and Pheidole morrisi [68] and Dolichoderus mariae [79] in the north of Florida. However, it is impossible to decide without special experiments whether these subtropical ants possess a real diapause, or they are quasi-heterodynamic and stop their development during the cold season due to direct influence of low temperature.
5. Two seasonal strategies of brood rearing in heterodynamic ants
Analyzing the structural diversity of heterodynamic seasonal cycles in ants, we identified two fundamentally different directions in their evolution, and accordingly, two seasonal strategies for brood rearing [18].
5.1. The strategy of prolonged brood rearing
The ants are more likely to follow the strategy of prolonged brood rearing. This strategy is based on the ability of larvae to enter into a diapause and to continue development over the next summer (Figure 1). Depending on the composition of the overwintering brood and the stage at which diapause is observed; we distinguish two structural types of developmental cycles [80].
Figure 1.
The strategy of prolonged brood rearing. Further explanation in the text.
Aphaenogaster type. Larvae fall into a diapause at the end of summer and all the remaining pupae manage to complete the development before the onset of colds, but the queens have no diapause and do not cease oviposition until the late autumn. Therefore, not only diapause larvae but also eggs and young larvae overwinter and survive, at least partially, during the winter. Thus, the formation of the wintering population of the colony is determined by the appearance of larval diapause in species with this type of seasonal cycle. These species are apparently limited in their distribution by subtropics and the southernmost regions of the temperate zone.
Myrmica type. The diapause starts both in larvae and queens at the end of summer. Therefore, before the winter comes, larvae emerge from the laid eggs, and all pupae develop into adults. The overwintering brood is represented only by larvae. This annual cycles are typical for most ant species living in a temperate climate zone. The induction of diapause in both larvae and queens plays an equally important role in the synchronization of Myrmica type cycles with the annual rhythm of the climate and in the formation of the wintering composition of the colony.
The annual cycles of ants that overwinter with brood have the most complex seasonal structure (Figure 1). All hibernating larvae usually pupate during the summer season. They give the first peak in the number of pupae in the nests. As a rule, alate females and males develop from most of them. Some of the larvae that emerged from the eggs which were laid in spring and early summer can pupate during the same growing season. This is the so-called a rapid or summer brood, according to Brian [81, 82]. It develops without diapause and gives the second peak in the number of pupae. All other larvae that emerge from the eggs within the season, fall into a diapause, hibernate and finish their development only next summer. This is so-called a slow or winter brood [81, 82]. Thus, two complete cycles of brood development from the egg to imago take place in a colony during each year (Figure 1): a summer cycle that begins and ends within one growing season, and a winter cycle, in which larval development is interrupted by their diapause.
Rapid brood is found in most species from the temperate zone. It may be absent in species and populations from the northern regions, where summer is short, as well as in species with very slow individual development. For example, the rapid brood is absent in Amblyopone pallipes from Massachusetts [83]. It can be assumed that this species is characterized by very slow development, although there are no data on this problem. The Japanese ant Leptanilla japonica has an extremely specialized annual cycle without rapid brood [73]. The queen lays the only portion of eggs at the end of July to the beginning of August. The larvae develop, overwinter at the last instar and pupate in July of the following year. They develop very synchronously, so that at each moment there is only one ontogenetic stage in the nest. Adults emerge from pupae at a time when larvae appear from the next portion of eggs. Thus, in spite of the fact that this species lives in subtropical climate, only one cycle of brood development occurs during a year.
The strategy of prolonged brood rearing is extensive inherently, as it is realized by stretching of the development of individuals for two or more summer seasons. The appearance of larval ability to fall into a diapause gave the ants a unique way for adaptation to the life in a temperate climate and especially in high latitudes. Therefore, the strategy of prolonged development is most common among the ants living there. It has a number of adaptive advantages.
5.1.1. More complete use of a favorable period for the development and available thermal resources
Since the larvae are always in the nest, the workers can feed them from early spring to late autumn. Immediately after the end of the winter, as soon as it becomes a little warmer, the ants carry larvae from the underground chambers to the upper, warmed by the sun, horizons of the nest (beginning with the largest larvae), creating the best opportunity for larval growth and development. As it becomes warmer and the amount of available food increases, the ants carry more and more small larvae to the upper levels of the nest and begin to feed them. According to Peakin for Lasius flavus, the overwintered larvae of the first and second instars remain in the deep and cold nest chambers for the longest time and therefore complete their development only by the end of the summer [84]. Just the same was recorded for Temnothorax nylanderi by Plateaux [69, 85]. In the spring, the workers of this species also start with the feeding of the largest overwintered larvae which develop into alates.
The autumn period of larval rearing is also of great importance for most ants with a wintering brood. For example, in the central part of European Russia in the colonies of M. rubra, the pupation of larvae ceases, as a rule, in the first half of August due to the onset of larval diapause. However, until the middle of September, the larvae continue to hatch from remaining eggs, to grow and develop. New third instar larvae appear and soon fall into a diapause. Moreover, in Myrmica diapausing, third instar larvae retain their ability to feed and to grow slowly (without the development of imaginal buds), and therefore, they continue to gain the weight in the autumn period [9, 86]. The same autumn growth of the last instar larvae was observed in Leptothorax species [69]. According to our data, larval growth in diapause state is a characteristic feature primarily for species in which larvae enter into a diapause and overwinter in the last or all instars, Aphaenogaster, Lasius, Leptothorax, Manica, Messor, Myrmica, Solenopsis, Temnothorax and Tetramorium and a number of species in genera Camponotus, Crematogaster and Monomorium.
Thus, in the species using the strategy of prolonged brood rearing, workers are engaged in feeding of larvae until the final onset of cold weather and give them all the surplus food produced during this period (minus the amount of nutrients that the workers accumulate in their fat body). This makes it possible to maximize the total mass of the wintering brood and, consequently, to grow up earlier the first workers, as well as reproductives, next spring. In addition, the biomass accumulated by larvae is also a reserve of nutrients for the colony: in the case of food shortage, ants can eat a part of the brood (mainly eggs and small larvae) in order to survive and to feed the largest larvae [2, 87].
5.1.2. The ability to adapt to the duration of the warm period of a year by changing the amount of the rapid brood
Such adaptation path can be realized during the penetration of the ants into more northern areas [88] and in connection with the local variability of climatic conditions from year to year. In the north, where summer is short and heat resources are limited, the ants can grow up much less number of the rapid brood than in the south. For example, in the south of France, Temnothorax unifasciatus has numerous rapid brood, and in cooler Belgium, only small amount of it [89]. According to our data for Lasius niger, L. flavus, Myrmica rubra, M. ruginodis and M. scabrinodis from the central part of European Russia, all overwintered larvae pupate in the spring and in the first half of summer, and then a numerous rapid brood larvae also pupate. Simultaneously, at the latitude of St. Petersburg, where the warm resources available to the ants of these species are much less, usually only a small part of larvae emerging from the eggs pupate during the same summer.
In the far north, where the summer is even shorter and the warmth is even less than in St. Petersburg, the ants generally never have rapid brood [88]. This was demonstrated in our studies on Leptothorax acervorum and Myrmica kamtschatica from the upper reaches of the Kolyma River and M. rubra and M. ruginodis from the coast of the White Sea near the Arctic Circle. Similar changes in the amount of rapid brood occur when the average summer temperature increases or decreases from year to year. For example, in cool summer, Temnothorax nylanderi may have no rapid brood, although it is usually quite numerous [69]. We observed the same in St. Petersburg region for Lasius niger, Leptothorax acervorum, Myrmica rubra, M. ruginodis and M. scabrinodis, in colonies of which there was no rapid brood in cool years, and even some larvae could stay for repeated overwintering.
5.1.3. The ability to stretch the development of larvae for two or even three summer seasons
Lack of thermal and/or nutritional resources during the summer is not uncommon situation in areas with cold temperate climate. As a result, some overwintered larvae that do not reach the size sufficient for pupation, fall into a diapause repeatedly and hibernate the second time. Repeated overwintering of some larvae was noted for Camponotus aethiops [90], Temnothorax nylanderi [69], Leptothorax acervorum [91], T. grouvellei [92] and Myrmica rubra [93]. We observed this phenomenon in our experiments with Aphaenogaster sinensis, Camponotus herculeanus, C. japonicus, C. aethiops, Lasius niger, L. flavus, Leptothorax acervorum, Manica rubida, Myrmica rubra and M. ruginodis. The possibility of repeated larval hibernation is of particular importance for ants living in the far north with an extremely short summer. According to our data for Myrmica kamtschatica and Leptothorax acervorum from the upper reaches of the Kolyma River, all wintering larvae pupate in warm years, but only a part of them if the summer is cold.
However, the number of larvae repeatedly overwintering probably cannot be significant in the colony. This is hampered by some social factors that limit possible changes in the structure of the seasonal developmental cycle when ants penetrate into more northern regions [88]. Therefore, ants never go over to the opportunistic strategy of stretching development for several years, so typical for many boreal and arctic insects [94, 95]. This feature restricts further spread of ants to high latitudes.
5.2. The strategy of concentrated brood rearing (Formica type)
This strategy presumes the obligatory completion of the development of larvae emerging from the eggs during one summer season, i.e. is typical for species that hibernate without brood (Figure 2). We named such annual cycles as Formica type [80], since they were first described for ants of this genus [7]. Diapause in larvae is absent, but it arises in queens at the end of summer long before the autumn cooling. Therefore, even the eggs laid by the latter have time to complete the development. All adults emerge from the pupae before cold weather comes on, and the ants prepare to overwinter. We can say that all brood is rapid in these ants.
Figure 2.
The strategy of concentrated brood rearing. Further explanation in the text.
After the onset of queen diapause, new eggs stop to appear and all existing brood gradually completes development. The diapause of queens should not occur too early, otherwise the period available for brood development would be actually reduced. Simultaneously, if diapause arises too late in the season, many larvae and pupae would be caught by the onset of winter and destroyed by the cold. That is why the moment when queens enter into a diapause is the most important for the Formica type annual cycles. From the point of view of using available heat resources, the strategy of concentrated brood rearing is less effective than the strategy of prolonged development. It can be realized in a temperate climate zone, where summer is short, only in the combination with increased brood developmental rate, i.e. by intensifying developmental processes, which becomes extremely important for northern species and populations living in areas with a particularly short summer.
Our studies have shown [42, 96] that this is really so: in Cataglyphis, and especially in Formica, individual developmental rates are significantly higher than in most species with hibernating larvae. Among the northern ants, Formica species have the shortest developmental times and develop almost twice as fast as Myrmica and Leptothorax. All six Formica species which we have studied were very similar in the duration of ontogenesis and the temperature sensitivity of the development [42]. Moreover, the development of Formica is much more thermal sensitive than in Myrmica, due to relatively higher temperature thresholds and a higher coefficient of linear regression of the developmental rate on temperature, which allows Formica ants to rear brood especially intensively at higher temperatures [42, 96].
According to our data, at temperatures of 25–26°C close to optimum temperatures for Formica species, their developmental time from the egg to the pupa is only 20–25 days, while it is 34–35 days in M. rubra. Moreover, at temperatures optimal for M. rubra (about 22° C), their developmental time from the egg to the pupa is 40–45 days, i.e. almost twice as much as Formica species. Thus, in Formica, brood rearing really occurs more intensively. In concordance with our observations for Serviformica species, in European Russia, their development from the egg to the adult can be consistently realized two or three times over the summer, i.e. these ants can rear two or three large batches of the brood. In Myrmica species, only one full cycle of development, rapid brood, passes in the colony during the summer season. The larvae of the second cycle enter into a diapause, overwinter and finish development next summer.
Thus, it can be assumed that raised rate of ontogenesis in Formica species completely compensates for the shortcomings of the strategy of concentrated development and allows to significantly increase the total amount of brood reared in the temperate climate during the summer. So far, however, it is difficult to say whether such a high rate of ontogenetic processes is a special adaptation that appeared in evolution during the formation of annual cycles of Formica type, or the strategy of concentrated development could arise only in ants that already had a high rate of development as preadaptational.
6. Review of heterodynamic annual cycles in ants
6.1. Seasonal cycles of Aphaenogaster type
For the first time, the seasonal cycle of ants of the genus Aphaenogaster was described in the work of Headley [5], who counted the quantitative composition of 46 colonies of A. fulva aquia (= rudis) in Ohio and recorded that eggs and larvae of all sizes remained in the nests for hibernation. These data were confirmed by Talbot, who conducted research on A. fulva and A. rudis in the more southerly state of Missouri [58, 97] and much later by Mizutani and Imamura on A. japonica in Sapporo (Japan) [98].
We found and investigated this annual developmental cycle on A. sinensis (Southern Primorye, Russia), A. gibbosa (southwestern Turkmenistan) and A. subterranea (Crimea) and suggested to separate it into a particular type [80]. Apparently, it is typical for the whole genus Aphaenogaster. In three species studied by us, a diapause of larvae of the last (third) instar arises at the end of summer. However, queens continue to lay eggs until late autumn, thus they do not have diapause. Therefore, eggs and larvae of all three instars stay in the nests for overwintering. At least some of the eggs and junior larvae hibernate successfully, both in the laboratory and in nature. We confirmed this fact during the excavation of nests in the south of Primorye and in the Western Kopet Dag (Turkmenistan) in early spring. In accordance with our data, the same seasonal cycle is typical for Tapinoma erraticum, T. karavaievi and, possibly, for some Messor species in Turkmenistan, as well as for Temnothorax species, although, in all these species, eggs cannot so successfully overwinter.
Thus, of all the currently known ants from the temperate climate going on hibernation with eggs, they probably overwinter happily only in the Aphaenogaster species. The incapacity of the eggs for hibernation is an obvious consequence of the impossibility of diapause onset at this ontogenetic stage in ants. Our experiments with the colonies of Lepisiota semenovi, Lasius niger, Myrmica rubra, M. ruginodis and Plagiolepis compressus, which we placed in summer into a refrigerator with a temperature of 3–5°C, have shown that the eggs of these ants died at low temperatures for 2 to 3 weeks and, of course, could not survive for a fairly long winter in the natural habitat.
Despite this, in subtropics and in areas with a very warm temperate climate, where winters are mild and short, seasonal cycles of Aphaenogaster type, which are characterized by the absence of queen diapause and overwintering with eggs and larvae of all instars, are likely to be widespread among ants. This is confirmed by some data. Thus, living in subtropics Linepithema humile [56], Polyrhachis vicina [77, 78], Solenopsis invicta, S. richteri, Rhytidoponera impressa [29] and R. metallica [99], always overwinters with eggs and all instar larvae. The presence of eggs and larvae of all instars in the nests during the winter and even the oviposition of queens at low autumn and winter temperatures are noted for Leptothorax niger, L. raeless and Temnothorax recedens in the south of France in a very warm temperate climate, almost subtropical climate [100]. Therefore, it seems obvious that the heterodynamic annual cycles of Aphaenogaster type are of subtropical origin. Some species can later penetrate into areas with a warm temperate climate, while retaining the “subtropical” structure of their cycle. However, in this case, eggs perish during the overwintering period due to more severe winter conditions.
Most species of the genus Aphaenogaster, including those studied by Headley [5] and Talbot [58, 97], are also confined to tropics and subtropics [2]. It can be assumed that some representatives of this genus, which have spread from subtropics to areas with colder and more prolonged winter, evolved some physiological mechanisms that increased the viability of eggs at low temperatures. To ascertain their nature, special investigations are needed. Thus, according to the literature and own data, in addition to Aphaenogaster species [5, 58, 80, 97, 98], the same seasonal cycle is typical also for some species of Leptothorax, Messor, Polyrhachis, Rhy-tidoponera, Tapinoma and Temnothorax, inhabiting areas with a subtropical and warm temperate climate.
6.2. Seasonal cycles of Myrmica type
The first studies of annual cycles of such type were fulfilled in the USA by Headley on two Leptothorax species [4] and by Talbot on M. schencki emeryana [6] and on two Temnothorax species [101]. Then, Passera studied in detail the seasonal development of Plagiolepis pygmaea in Southern France [102, 103], and Sanders investigated three Camponotus species in the south of Canada [104, 105]. The seasonal cycles of Messor capitatus [106], Camponotus vagus [107] and C. aethiops [108] in the south of France, Myrmica rubra in Belgium [109] and Paratrechina flavipes in Japan [110] were studied in the same detail.
The first study, in which the annual cycle of development was observed in the laboratory, belongs to Brian [81]. He maintained two colonies of M. rubra and M. ruginodis, in artificial conditions closest to natural. Later, he investigated the growth and development in several colonies of the same species, founded by females, before the production of alates [93]. The developmental cycle of Plagiolepis pygmaea [103] and a number of Leptothorax species [69, 75, 85, 89, 92, 111, 112] were studied in much greater detail. Plateaux kept the colonies founded by the inseminated females of Leptothorax under conditions simulating the change of seasons, from the moment of the colony foundation until the queen death and the natural extinction of the colony (5–12 years), i.e. recorded the complete ontogeny of all colonies [75, 92, 111, 112]. Similar studies have been conducted on Messor syriacus [113], M. incorruptus [114] and M. barbarus [115, 116]. Based on literature data, we can include in Myrmica type, the seasonal developmental cycle of European species Messor harvester [117] and Dolichoderus mariae [79] from the north of Florida (USA).
The larval stages, on which the diapause can occur, are extremely variable among ants with Myrmica and Aphaenogaster types of annual cycles. Five groups can be distinguished according to the instar composition of overwintering larvae [18].
6.2.1. Diapause in early (1st–3rd of 5 or 6) instars (Lepisiota, Plagiolepis, Tapinoma and some Camponotus)
Larvae of the first three instars hibernate in Plagiolepis pygmaea according to Passera [102, 103]. Our data for P. calvus, P. compressus, P. karawajewi and P. vladileni are completely consistent with this conclusion. In Tapinoma erraticum from France, most of the larvae hibernate at the first instar and only a few at the second and third [118]. We found the same in T. erraticum and T. karavaievi from Turkmenistan. We also observed the hibernation of the first and second instar larvae in Lepisiota semenovi from the same place.
Diapause in early instars is also typical for many Camponotus species. Larvae of European C. vagus always hibernate in the first instar [107]. Mintzer observed the same in incipient colonies of seven American Camponotus species (C. clariothorax, C. festinatus, C. laevigatus, C. modoc, C. planatus, C. rasilis and C. vicinus) under the laboratory conditions [119].
6.2.2. Diapause in middle (2nd–4th of 5–6) instars (Camponotus s. str.)
Sanders indicated that in C. herculeanus, C. noveboraceus and C. pennsylvanicus, living inside tree trunks in southern Canada, hibernating larvae could be divided into two distinct size classes, i.e. they were clearly at different instars, unlike the Camponotus species listed above [104, 105]. We observed the hibernation of larvae of the second, third and in a small number of the fourth instar in our experiments on C. herculeanus, C. japonicus and C. ligniperda, all belonging to the same subgenus Camponotus s. str. The first instar larvae never overwintered. In the natural nests of C. herculeanus in early spring, we also found mainly larvae of the second and third instar, but did not find larvae of the first and fifth instars at all.
6.2.3. Diapause in the late (3rd–4th of 4) instars (Harpagoxenus, Leptothorax, Temnothorax and Messor)
According to the literature data, the larvae of Messor incorruptus hibernate in the last two instars [114]. Delage noted that among the overwintering brood of M. capitatus there were no larvae of the first instar [106]. In the laboratory, we observed the hibernation of M. denticulatus, M. intermedius, M. subgracilinodis and M. structor and found that their larvae overwintered in the last (third and fourth) instars. In the nests of M. denticulatus and M. structor from Turkmenistan in the early spring, we always found only larvae of the last two instars.
Our investigations demonstrated that in Leptothorax and Harpagoxenus sublaevis larvae hibernated in the third and fourth instars, and larvae of the latter instar were clearly predominant among them. Only very rarely single larvae of the second instar could be found in overwintering colonies of these species. Larvae of Temnothorax may hibernate at all three instars, but larvae of the last instar obviously predominate among overwintering ones, for example, in T. lichtensteini, T. nylanderi, T. parvulus [69, 112], T. grouvellei [92], T. melas [75] and T. monjanzei [74]. We observed the same in T. unifasciatus and T. tuberum.
6.2.4. Diapause in the last (usually 3rd) instar (Manica, Diplorhoptrum, Leptanilla, Monomorium, Myrmica, Tetramorium)
Only larvae of the third (last) instar hibernate in all Myrmica species studied [109, 120, 121, 122]. Our studies on M. bessarabica, M. kamtschatica, M. lacustris, M. rubra, M. ruginodis, M. sabuleti, M. scabrinodis and M. transsibirica confirmed this. Larvae overwinter in the latter instar also in Leptanilla japonica, which inhabit the subtropical regions of Japan [73]. Our research work allowed us to supplement this group with the following species: Monomorium gracillimum, M. ruzskyi, Manica rubida, Solenopsis celatum, S. fugax and all Tetramorium species.
Larvae of the ants, hibernating at the last instar, are for the most part far from the completion of development, i.e. they are at the beginning or in the middle of this stage. To achieve the size required for pupation, they usually need a fairly long period of growth after overwintering. This fully applies to the largest of overwintering larvae that develop in spring into alate reproductives. An exception to this rule is Leptothorax acervorum and probably other species of the same subgenus, in which many of the larvae of reproductives almost complete their development before the winter.
6.2.5. Diapause in all (of 3–6) instars (sometimes except the first instar) (Aphaenogaster, Crematogaster, Lasius, Paratrechina, Camponotus, Tanaemyrmex)
This group includes Lasius flavus [84], L. alienus and L. niger (own data), Paratrechina flavipes [110], Crematogaster bogojawlenskii (own data) and some species of Camponotus. An exact study of the instar composition of overwintering larvae of C. aethiops in France showed that 85% of them were larvae of the first and second instars, 5% of the third to fourth and 10–15% of the fifth instar [108]. The results of our experiments and spring excavation of nests in nature confirmed that the overwintering brood of this species was represented by larvae of all instars except the first, with the obvious dominance of the second instar. We have shown that larvae of C. bactrianus and C. xerxes, species of the subgenus Tanaemyrmex as well, also hibernated in all instars except the first. Perhaps this feature is typical for the entire subgenus.
6.3. Seasonal cycles of the Formica type
According to the literature data and own observations, this annual cycle is typical for the ants of the entire tribe Formicini (genera Alloformica, Cataglyphis, Formica, Proformica, Rossomyrmex). This fact was firstly determined by Holmquist on Formica ulkei [123, 124] and later repeatedly confirmed by other researchers for same [125] and other Formica species: F. fusca [7], F. haemorrhoidalis [126], F. japonica [127] and F. yessensis [128]. Annual developmental cycles of ants of the subgenus Formica s. str. have been studied in detail by many authors (for example, see [129, 130, 131, 132]). There are also detailed experimental data for Cataglyphis cursor [133, 134, 135]. Many species of genera Alloformica, Cataglyphis, Formica and Proformica have been studied in Central Asia by Dlusskii [63, 136]. The seasonal cycle of the slave-maker ant Rossomyrmex proformicarum in the deserts of Semirechie (Republic of Kazakhstan) was described by Marikovskii [137]. Our experiments and field observations made it possible to add to this list Cataglyphis aenescens, C. emeryi, C. nodus, C. pallida, Formica cinerea, F. clara, F. cunicularia, F. gagatoides, F. lemani, F. picea, F pratensis and Proformica epinotalis.
Outside the tribe Formicini, annual cycles without wintering brood were found in Dolichoderus plagiatus, D. pustulatus [138] and D. quadripunctatus [139, 140] and in American harvester ants Pogonomyrmex occidentalis and P. montanus [59, 141, 142]. Prenolepis impairs from the north of Florida also overwinters without brood but has completely special annual cycle [60, 62]. These ants are active outside the nests and foraging from November to March, i.e. during the winter months; the rest of a year the nests are closed. During the foraging period, workers inside the nest accumulate reserve nutrients in the fat body and gradually become physogastric. In spring, their mass exceeds the normal by 2–3 times. During spring and summer, the ants stay in the nests in inactive state. In September, the queens lay a large number of eggs, and then their ovaries again become dysfunctional. The ants feed the larvae emerging from eggs only at the expense of fat stocks of physogastric workers. Both workers and winged reproductives grow up from this single batch of brood. By the time of the resumption of foraging, there is no brood in the nests.
We found the annual cycle of Formica type in Ponera coarctata from the southern coast of the Crimea (Karadag). Investigation of several dozen nests in early September demonstrated that brood, except for a small number of pupae, was no longer present in them. In the colonies transferred from natural nests to the laboratory, the eggs did not appear even when the ants were kept at a long day photoperiod and at temperatures of 25–28°C for 2 months. Thus, the reproductive diapause of queens in this species was as stable as that of Formica ants. According to Talbot, Ponera pennsylvanica in Missouri (subtropics) also hibernated without brood [58]. Similar seasonal cycle was described for another Ponerine ant, Odontomachus brunneus from the north of Florida (subtropics) [143]. This species spend half of a year (from November up to April) without any brood in the nests. This allows us to assume that queens of Odontomachus brunneus have winter reproductive diapause.
7. Two types of the regulation of heterodynamic annual cycles
The main characteristic feature of homodynamic and quasi-heterodynamic types of annual cycles is a purely exogenous control of the development, and the key factor is environmental temperature. Heterodynamic species have a much more complex regulation of seasonal development, usually based on a combination of exogenous and endogenous mechanisms. We divided all heterodynamic ants into two groups, which differ substantially in the principles of the regulation of the annual cycle, exogenous-heterodynamic species and endogenous-heterodynamic species [18].
The first of them is characterized by the possibility of continuous and unlimited development under optimal conditions. The diapause is optional and occurs only when the temperature is lowered. Such annual cycles we call exogenous-heterodynamic. They are typical for all species of the genera Messor, Monomorium, Solenopsis and Tetramorium we have studied and also for Camponotus xerxes and Tapinoma karavaievi. At temperatures above 25°C, as well as at diurnal thermal periods 20/30°C, they all behave like true homodynamic species. Under these conditions, we observed continuous development without any signs of deterioration in experimental colonies for two or more years [144]. At the same time, at temperatures below 23–25°C, the egg laying and the larval pupation soon stop, or only pupation (in some species of Messor, which can hibernate with eggs), i.e. development finishes under the influence of suboptimal temperatures. If the temperature is then raised, the development will resume after a while. However, it can again be blocked by lowering the temperature, and once again stimulated by its increase. Similar experiments can be repeated with the same colony of ants many times and oft with the same result [144].
Thus, exogenous-heterodynamic species are distinguished by facultative winter diapause in larvae and queens. Developmental delays in a colony are purely exogenous and ensue as a straight reaction to the influence of external environmental factors, primarily, of temperature when it becomes not optimal for the development. Moreover, inhibition of development is unstable and easily disrupted when the temperature rises. However, these developmental delays are not just a consequence of cold coma but, namely, are the form of diapause (for more details, see [18]). This is not a state of elementary quiescence as in species with quasi-heterodynamic annual cycles, because this kind of diapause starts when environmental temperature still significantly exceeds the developmental threshold. Other essential feature is that the diapause arises not directly after the temperature decline but with some lag. Additionally, this diapause is invertible and may be terminated or induced again several times by consistent rising or decreasing the temperature, but each time after a little delay. The second important property of exogenous-heterodynamic species is the distinct change of their reaction to temperature during overwintering as a result of cold reactivation. After natural overwintering or after exposure in a refrigerator to 3–5°C during 2–3 months, the development and pupation of larvae recommenced and proceeded for a long period at 20°C and even at 18°C (in some Tetramorium) in all species studied [144]. This difference in the reaction to temperature is another indication of the existence of diapause in exogenous-heterodynamic species.
Most of the ants from temperate zone belong to the second group of species which is characterized by endogenous-heterodynamic annual cycles. The diapause arises due to internal factors (endogenous timer) and no external conditions can prevent it [18, 145]. Even under long day conditions and optimal temperatures, including the diurnal thermal periods, which are the most favorable thermal conditions for ants [146, 147, 148], the development in colonies of these species necessarily ceases, and the phase of dormancy in the annual cycle begins.
Thus, the diapause of endogenous-heterodynamic species is obligatory for the colony which has an internally limited intrinsic seasonal cycle of brood rearing. External environmental factors, such as temperature and photoperiod, also participate in the regulation of annual cycles of these species, playing a corrective role, i.e. in varying degrees adjusting the duration of the cycle to the climatic peculiarities of a particular summer season. But the regulation of the cycle is still based on processes that are endogenous for the colony [18, 145]. According to our data, the following species living in temperate climate belong to the group with endogenous-heterodynamic annual cycles: Aphaenogaster, Camponotus, Cataglyphis, Crematogaster, Formica, Harpagoxenus, Lasius, Lepisiota, Leptothorax, Manica, Myrmica, Plagiolepis, Ponera, Proformica and Tapinoma [18, 80, 96, 145, 146, 147, 149, 150, 151]. Analysis of the literature data from our conceptual positions allows us to classify the following species as endogenous-heterodynamic: Aphaenogaster subterranea [152], Camponotus herculeanus, C. ligniperda, C. noveboraceus, C. pennsylvanicus, C. vagus [105, 107, 153], Cataglyphis cursor [133, 134, 135], Crematogaster scutellaris [154, 155, 156, 157], Formica sanguinea [158], F. ulkei [159], the genera Leptothorax and Temnothorax (with the exception of subtropical ones) [69, 75, 85, 91, 112], Myrmica rubra [82, 160, 161, 162], Plagiolepis pygmaea [103] and Odontomachus brunneus [143].
The gradual decrease of a colony capability to produce new eggs and to grow up non-diapausing larvae takes place during the summer season. At the same time, there is the increase of the bias for diapause as a consequence of the ongoing endogenous physiological and social processes. Moreover, the ant colony gradually acquires the sensitivity to the day length (to the photoperiod). The increasing photoperiodic sensitivity of a colony strongly changes the reaction to temperature. Because of these processes, at the end of the summer, the decrease of temperatures and the shortening of the day length (in some species only) contribute to the onset of diapause, thereby reducing the period of egg laying and larval development. The same impact of external factors to the colony’s life cycle was found in all ant species that we studied [80, 96, 146, 163, 164, 165]. So, the duration of a colony’s annual cycle of brood rearing in nature is controlled both by an endogenous timer and by exogenous environmental cues, such as temperature and photoperiod (in some species). These environmental conditions adjust the date of diapause onset to the climatic features of a given year.
Temperature control of diapause is the most universal in ants. The higher temperatures delay the onset of diapause both in larvae and adults, whereas the lower temperatures always accelerate the process of the beginning both in larvae and adults. On the contrary, photoperiodic control of diapause is unexpectedly uncommon among ants. For the first time, the photoperiodic responses were revealed in M. rubra and M. ruginodis [163]. It has been shown that the diapause in larvae and queen appeared more quickly when the day length decreased in the range from 16 to 13 h [163, 164]. The diapause both in queens and larvae ceased when inactive ant colonies in autumn condition were exposed to day lengths of 15 h [166]. Later on, Hand [167] and Brian [168] confirmed the presence of the reaction to photoperiodic conditions in M. rubra.
The subsequent extensive investigations nonetheless have shown that apart from Myrmica only few ant species used photoperiod as an ecological cue which controls all life processes in a colony: development, oviposition and diapause onset. We have found that photoperiodic conditions affected the induction of diapause only in Aphaenogaster sinensis [80] and Lepisiota semenovi [169]. Additionally, we have observed higher incidence of diapause in larvae under short-day conditions than in larvae under long-day conditions in Camponotus herculeanus, Leptothorax acervorum and Manica rubida. Thus, the genus Myrmica represents a rather curious exception among temperate ants since all Myrmica species studied so far possess clear-cut photoperiodic responses, which control the induction and termination of diapause [149, 150, 163, 164, 170, 171].
Thereby, the seasonal development of most ant species from temperate climate regions depends on inner timer in combination with environmental temperature, which triggers the onset of diapause. The environmental factors can modify the duration of the seasonal brood-rearing cycle within broad range in most species of the genera Aphaenogaster, Crematogaster, Lasius, Myrmica and Tapinoma. For example, suboptimal temperatures of 17–20°C and short days vastly bring closer the beginning of diapause in larvae and queens of Myrmica rubra and M. ruginodis [163]. Whereas at long days and a temperature of 25°C, egg laying and the development and pupation of rapid brood larvae continue for several months without interruption [149, 164].
The seasonal cycle of oviposition and development in other species is controlled predominantly by the endogenous mechanisms. In these ants, the moment of diapause onset depends only slightly on environmental conditions. Temperature hardly modifies the intrinsic length of the queens’ oviposition period in all studied Formica species [96, 165]. The annual brood-rearing cycle in Cataglyphis species and the species from subgenera Camponotus s. str. and Leptothorax s. str. is also relatively independent of the environment.
8. The regulation of diapause in larvae
The diapause of larvae in ants is facultative in most case, i.e. a given larva can either develop directly or fall into a diapause depending on the circumstances. Temperature can affect larval development and induce diapause both directly and through the nurse workers. Detailed studies performed on M. rubra have made it possible to prove that ant larvae themselves were not able to sensate the photoperiodic conditions in which they were placed [172, 173]. Actually, the development of larvae was under the influence of workers whose physiological state and behavior strongly depended on the day length because they received the information about the photoperiodic conditions from the external environment. In laboratory experiments, we easily obtained the workers and larvae in alternative physiological states either by keeping the ant colonies under long-day conditions from the spring and by activating them with long-day photoperiods in autumn (getting of physiologically active, non-diapause individuals) or by exposing the ant colonies to the influence of short-day conditions during 1 month (getting of physiologically inactive, diapausing individuals). We have shown that non-diapause workers induced growth and pupation of summer larvae and interrupted the diapause of autumn larvae, while diapausing workers were incapable to support a high growth rate of larvae and forced them to fall into a diapause [13, 170]. We examined the characteristic features of social control by workers over larval development in several ant species and created experiments using workers and larvae in alternative physiological states. The experimental scheme included four sets: (1) spring state: physiologically active workers (after hibernation) with physiologically active larvae; (2) termination of larval diapause: physiologically active workers with diapausing larvae; (3) induction of larval diapause: diapausing workers with physiologically active larvae and (4) autumn state: diapausing workers with diapausing larvae.
The results of these experiments demonstrated that ant species studied fundamentally distinguished from each other [13, 14, 15, 16, 170, 174]. In Camponotus herculeanus, C. japonicus and several Tetramorium species not overwintered physiologically inactive diapausing larvae, which were provided with food by spring physiologically active workers, developed rapidly and pupated within a short period, whereas overwintered larvae placed into the nests with autumn diapausing workers did not develop and pupate at all or only a few of them pupated sometimes. Thus, the workers of these ants exercise full control over the development and the diapause of their larvae. However, in Leptothorax acervorum, we found an entirely opposite situation: autumn workers could not prevent development of spring larvae and they all pupated. At the same time, overwintered workers stimulated the development of less than half of diapausing larvae. In Myrmica rubra, M. ruginodis, M. lobicornis and Lasius niger, we observed an intermediate situation: only some spring larvae pupated when fed by autumn workers, and also far from all autumn larvae finished the development under the care of spring workers.
Thus, the forms of social influence of workers on larval diapause are diverse in ants and range from nearly absolute control (in Camponotus and Tetramorium), when the physiological state of workers completely defines the fate of larvae, to rather weak effects when diapausing workers are unable to prevent the pupation of most of overwintered larvae, and spring workers are capable of inducing the development and pupation of only a few diapausing autumn larvae (in Leptothorax). In most species (Lasius, Myrmica), however, the intermediate variants of diapause control are realized. There is some evidence that Myrmica workers can manipulate the development of larvae via changing the intensity of tactile stimulation and the frequency of feedings [12, 13, 170, 174]. Probably, in the cases when larvae hibernate at younger instars (as in Camponotus s. str.) and need a long period of growth to complete development, diapausing workers cannot provide them with the necessary food. At the same time, spring workers are able to effectively stimulate the development of young diapausing larvae. On the other hand, large larvae, overwintered in the last instar (as in Leptothorax), can finish the development even without receiving enough food from diapausing workers. However, the diapause of these larvae is deeper and more durable, and the spring workers cannot break it.
In many ant species with endogenous- and exogenous-heterodynamic seasonal cycles, the larvae fall into a diapause in the last larval instar. These diapausing larvae continue to feed and to grow slowly and can attain a significantly larger size before overwintering. This larval growth in diapause state is very important for the process of caste differentiation in Myrmica [86]. Continuing to increase in the size the diapausing larvae in reality do not develop progressively, as far as the differentiation and enlargement of their imaginal buds do not occur. Only these well-grown large diapausing larvae potentially may become the female reproductives in spring [86]. Subsequently, Plateaux [69] described the identical phenomenon for L. nylanderi. In accordance with the results of our studies, slow growth of larvae in diapause state is an attribute of the ant species from the genera Camponotus (some species), Crematogaster, Lasius, Lepisiota, Leptothorax, Manica, Messor, Monomorium, Myrmica, Solenopsis and Tetramorium. The physiological nature and types of diapause in ants are examined in the paper of Kipyatkov [18].
9. Evolution of annual cycles of development in ants
Problems of the origin and evolution of the diapause and seasonal cycles of insect development attracted the attention of many researchers (for example, see [175, 176, 177]). The main conclusion reached by most authors is that the evolution of seasonal adaptations occurs largely beyond direct connection with the phylogeny of taxa, and all the elements and parameters of seasonal development known to us, including the diapause itself, the types of cycles, photoperiodic reactions and other regulatory mechanisms, repeatedly, independently and in a variety of ways arise in the evolution of insects. One and the same goal of adapting to certain seasonal environmental conditions can be reached in a variety of ways by combining a whole range of known (or even not yet described) physiological mechanisms [176, 177]. Such a clearly expressed diversity of evolutionary solutions makes it very difficult to analyze the possible ways of the evolution of seasonal development cycles, even within not very large groups of organisms. Nevertheless, some authors proposed various specific sequences of evolutionary events to explain the origin and development of photoperiodic reactions and other physiological mechanisms controlling the diapause and other seasonal adaptations [175]. The most promising approach to the problem of the evolution of seasonal adaptations can be the identification of basic adaptation syndromes, for example, structural types of the annual cycles, diapause forms, ways of synchronizing the cycle with the seasonal rhythm of the climate, etc. Then, it may be productive to look for correlations between all these adaptations and possible ties with the specific environmental conditions in which they are realized and also to analyze the occurrence of the phenomena under study within different taxa.
More and more data appear on the role of diapause in the regulation of the life cycles of insects in tropics. In these regions, the diapause does not always prove to be adapted specifically to the extreme conditions of the abiotic environment, but is often associated with seasonal variations in food availability, migration and reproduction processes. This makes it possible to speak with confidence about the tropical origin of the diapause in many insects [70, 176]. Moreover, the winter hibernation of insects is a relatively recent evolutionary acquisition that arose only after the formation of a glacial climate on the Earth [44]. The available facts do not completely exclude the hypothesis of the possible occurrence of the diapause by the gradual deepening of the developmental delays that first arise exogenously under the direct influence of cold, dryness, lack of food or other unfavorable conditions. The diapause is a fairly simple adaptation from the point of view of the possibility of forming its regulatory hormonal mechanisms, and therefore it could arise in evolution many times and in different ways [176, 177].
Turning to the analysis of the main trends in the evolution of the annual developmental cycles in ants, two most important and closely related questions should be pointed out. First, it is the origin of different forms of diapause, and secondly, possible ways of forming of endogenous-heterodynamic developmental cycles with obligate diapause. It should be borne in mind that the family Formicidae, in its evolutionary origin, is undoubtedly associated with the tropical regions of the Earth, where most of the species of ants live now. From tropical regions, these insects penetrated into zones with temperate climate, forming new species and taxa of higher rank [2, 63]. Another interesting issue, namely the evolution of the structure of the seasonal cycles in ants during their distribution to the north, has been discussed in detail in a special article of V. E. Kipyatkov [88].
It is possible to imagine at least two possible ways of the origin of heterodynamic annual cycles in ants living in temperate and cold climatic zones.
9.1. Subtropical (quasi-heterodynamic) path of evolution
We suppose the possibility of direct adaptation of homodynamic species which penetrate from the tropical regions into subtropics and further into the zone with temperate climate and with cold enough winter. In this case, they do not form a real diapause, and at first acquire the ability to overwinter in the state of a quiescent (cold coma) but suffer from more or less strong mortality, i.e. quasi-heterodynamic seasonal cycles appears. The diapause evolves later and seasonal development becomes exogenously heterodynamic.
The reality of this path of evolution is almost unquestionable. It has been illustrated above by a number of examples of quasi-heterodynamic seasonal cycles, in particular, by the example of two Pheidole species studied experimentally. Some of the quasi-heterodynamic species (Linepithema humile, Solenopsis invicta, S. richteri) are known to have penetrated recently into areas with subtropical and temperate climate. They represent a magnificent model of the evolution of heterodynamic cycles, but practically were not investigated experimentally. It is completely unclear; for example, do the populations of S. invicta that penetrated far enough to the north in the USA, shape some kind of a diapause, or in winter the development delays because of a quiescent, like in the populations from subtropics? The same question applies to the populations of L. humile in Southern France. An experimental study of these ants, as model systems, would make it possible to test the possibility of quite quickly (over several decades) occurrence of a diapause in quasi-heterodynamic species.
Probably, quasi-heterodynamic annual cycles with the brood death in late autumn arise in the evolution of many homodynamic ants, which penetrate from tropics into subtropics and further into regions with a warm temperate climate. The nomadic ants Neivamyrmex on the north of its distribution area in the USA live in regions with a fairly cold winter. Investigations were conducted on N. nigrescens by Schneirela in the southeast of Arizona at altitudes of about 1660 m above sea level [31]. He has shown that in the autumn, when the nights became cold, these ants with nocturnal activity ceased foraging activity. The lack of food compelled the queen to stop egg laying, the workers destroyed the remaining brood and adult ants hibernated in a shelter. In the spring when the temperature rose, the queen started oviposition. When larvae appeared, the ant colony gradually restored the cycle of nomadic behavior and brood rearing typical for the summer period [31]. Schneirela assumed the presence of direct influence of temperature on the development of these ants. However, this idea was not confirmed by experiments.
The second real way of verification of our assumption is to study the species taxonomically close to exogenous-heterodynamic ants from the zone with warm temperate climate, but inhabiting subtropics and tropics. The most promising in this respect are the genera Monomorium and especially Tetramorium. All of the species of these genera, which we studied, inhabit the temperate climate zone and are exogenous-heterodynamic.
If the onset of a diapause and exogenous-heterodynamic development on the basis of quasi-heterodynamic seasonal cycles seems quite plausible, then the possibility of further evolution in this direction toward the formation of endogenous-heterodynamic cycles with obligate diapause is far from obvious. We do not have at present any definitive evidence of the reality of such an evolution, but in our experiments we found distinct manifestations of endogenous regulation of development in most exogenous-heterodynamic species [144]. The possibilities of evolutionary transition from exogenous- to endogenous-heterodynamic development within the group of closely related species are indirectly confirmed by the following three examples.
Two closely related species of the genus Tapinoma from Turkmenistan were investigated in sufficient detail experimentally and under natural conditions. T. karavaievi is a species widespread in the plains of Central Asia. It does not rise to the mountains higher than 600 m above sea level, i.e. lives in a fairly warm temperate climate [136]. Almost in all parameters of the regulation of seasonal development, this species is exogenous-heterodynamic. In the most optimal temperature conditions, we observed almost non-stop development in its colonies. The diapause of larvae in T. karavaiev, as a rule, terminates when the temperature rises [144]. Nevertheless, the endogenous regulatory component was distinctly expressed in this species at lower temperatures, and under conditions of free temperature selection in the temperature gradient installation, we observed a smoothed spontaneous rhythm of development [145]. Another species studied, T. erraticum, is common in southern Europe and the Caucasus, and in Turkmenistan occurs only in the mountains, i.e. clearly gravitates toward a cooler climate than T. karavaievi [136]. Experiments have shown that T. erraticum belonged to the group of endogenous-heterodynamic species and had a very stable winter larval diapause, which could not be disturbed by a simple increase in temperature.
The second example is a pair of taxonomically related species of the genus Monomorium, M. kusnezovi and M. ruzskyi. Due to the existence of significant geographical variability in morphology for a long time, many researchers have combined these ants into one species under different names and confused them [136]. The gradual accumulation of collection materials made it possible to find out that everywhere on the plains of Central Asia there is one variegated in color species, M. kusnezovi. Its mountain populations differ from the desert ones not only in smaller sizes and dark colors, but also in some peculiarities of biology [63]. Later, large differences between the queens of these two forms were found and the mountain form was described as M. ruzskyi [136].
Our observations have shown that M. kusnezovi and M. ruzskyi often met together in the foothills and lowlands of the Kopetdag. However, despite the taxonomic proximity of these two species, their annual cycles differ significantly. M. kusnezovi is quasi-heterodynamic, i.e. the diapause of the brood and the reproductive diapause of queens are absent, so eggs and larvae of all ages remain in the nest during the winter and a significant part of them perish. At the same time, the annual cycle of M. ruzskyi is of Myrmica type, i.e. it is endogenous-heterodynamic and is characterized by a fairly stable diapause in larvae of the last (third) instar and in queens. Therefore, in the colonies of this species, only the larvae of the last instar hibernate, and they are always numerous. Thus, the features of the structure and regulation of the annual cycle make it possible to differentiate these closely related species no worse than the morphology of their queens.
We studied two Camponotus species, C. xerxes and C. aethiops, which belonged to the subgenus Tanaemyrmex, but differed dramatically in regulation of seasonal development. C. aethiops possesses an obligate, very stable diapause and endogenous-heterodynamic regulation of the annual cycle, while in C. xerxes there is an unstable diapause and mainly exogenous regulation. At a temperature of 25–27°C, we observed continuous development in the colonies of C. xerxes for more than 2 years. These differences are obviously due to the fact that C. aethiops lives in a colder climate (Central and Southern Europe, Crimea, Caucasus and mountains of Central Asia) than C. xerxes (the desert plains of Afghanistan, Iran and Turkmenistan).
9.2. Tropical (preadaptational) path of evolution
Extremely few examples of heterodynamic annual cycles in tropical ants allow us to assume that such species, already possessing diapause (preadaptation), could probably easily penetrate into subtropics and further into the temperate climate zone, using the ability to form a diapause for experiencing a cold winter. The causes for diapause emergence in tropical ants can be diverse. For example, it can be the necessity for survival during the arid or excessively wet seasons of a year, during the periods of shortage or inability to obtain food, etc. It is also possible that the ability to diapause arose as a way of solving internal problems for the colony related to the regulation of development, the processes of caste differentiation and reproduction. Moreover, in tropics, both exogenous and endogenous mechanisms of diapause regulation could be formed, which under the new conditions of a temperate climate, and could be used to synchronize the onset of a diapause with the beginning of a cold season of a year.
Unfortunately, this scheme is almost entirely speculative, first of all, because tropical species with heterodynamic annual cycles have not been investigated experimentally so far, and we know absolutely nothing about the prevalence and nature of diapause in tropical ants. The only argument indirectly confirming the possibility of the tropical origin of heterodynamic annual development in many ants is the rather wide prevalence of endogenous-heterodynamic cycles with obligate diapause among ants inhabiting subtropical and warm temperate climate zone. Among the 39 species of ants from Turkmenistan and the southern coast of the Crimea which we experimentally studied, 18 (46%) are endogenous-heterodynamic. An analysis of a few works, in which at least the simplest laboratory experiments were carried out, allows us to classify as endogenous-heterodynamic four more species from the zone with warm temperate and subtropical climate: Plagiolepis pygmaea [103], Camponotus vagus [107], Temnothorax monjanzei [74] and T. melas [75]. Are we correct to assert that in all these ants, the predominantly endogenous regulation of the annual cycle with the obligate diapause has evolved here in areas with a very warm or subtropical climate, or is it wiser to think that it originated in tropical ancestors of some of these species at least? The answer to this question is currently not possible due to lack of sufficient evidence.
9.3. Taxonomic position and seasonal adaptations
An example of close relationship between taxonomic position and seasonal adaptations is the Formicini tribe. All species studied use the strategy of concentrated brood rearing and are endogenous-heterodynamic with the obligate and very stable diapause of queens and workers and the apparent dominance of endogenous regulatory mechanisms. Probably, such a system of seasonal adaptations evolved already in the ancestors of this tribe.
All studied species of the genus Myrmica have a very similar structure of the seasonal development cycle [178, 179] and, according to our experiments, have photoperiodic regulation of a diapause rarely seen in other ants [18]. We have experimentally investigated the seasonal development of three species of the subgenus Camponotus s. str., C. herculeanus, C. japonicus and C. ligniperda and found almost complete similarity between them in all respects: endogenous-heterodynamic regulation, obligate and very stable diapause of queens and larvae, the same overwintering stages (second instar larvae) and, finally, absolute control by workers over the onset and cessation of larval diapause [16]. According to available literature data, C. vagus from the same subgenus has identical characteristics of the seasonal cycle [107]. Finally, all species of the genus Tetramorium, which we studied, are distinguished by exogenous-heterodynamic regulation of seasonal development and temperature-unstable diapause.
Other examples relate to fairly definite correlations between taxonomic position and overwintering stages in a number of ants genera. In all species of Leptothorax and Messor, larvae diapause and overwinter mainly in the last two instars; in Myrmica and Tetramorium, in the last instar, obligatory; in Plagiolepis, in the first three instars and in Lasius, in all instars. Finally, all Aphaenogaster species studied so far do not have a diapause of queens and therefore hibernate with eggs and larvae in all instars. At the same time, directly opposite situations are known, when closely related ants have completely different seasonal adaptations. They are already mentioned above when comparing the following pairs of species: T. karavaievi–T. erraticum, M. kusnezovi–M. ruzskyi and C. xerxes–C. aethiops.
Thus, it can be argued that in the evolution of ants, correlations could arise between the nature of seasonal adaptations and the phylogeny of taxa, but no less common are the cases of the absence of such connections, i.e. the appearance of significant differences in the structure and regulation of the annual cycle between related species and, on the contrary, the parallel and independent formation of very similar adaptations in different phylogenetic branches.
10. Conclusions
Most tropical ants demonstrate homodynamic development. They do not exhibit any developmental delays and all-year round the ontogenetic stages from egg to pupa exist in their nests. Some of the quasi-heterodynamic species have permeated into the regions with warm temperate climate but a true diapause did not evolve. In these species, the brood development stops only at temperatures falling below the developmental threshold (consecutive dormancy). So, the ants spend the winter in the state of a quiescent (cold coma), while more or less high mortality rates are observed in their colonies. Most temperate and all boreal climate ants are true heterodynamic. They manifest a true deep winter diapause (prospective dormancy) in their annual cycle.
Heterodynamic ants use two main seasonal strategies with respect to brood rearing. The ants are more likely to follow the strategy of prolonged brood rearing. It is distinguished by the following features: (1) larval diapause is facultative and controlled by environmental (temperature, photoperiod) and social (worker care, queen influence, pheromones, etc.) factors; (2) only some larvae develop from egg to pupa within the same summer season without overwintering (this rapid brood, or summer brood, yields only workers); (3) a large proportion of larvae delay their development, continue to grow in autumn, overwinter in diapause and pupate the next summer (this slow brood, or winter brood, yields both workers and alates).
The strategy of concentrated brood rearing is distinguished by the following features: (1) larvae have no dormancy and complete their development during the summer; (2) the development of all brood stages is thus restricted to the growing season; (3) only queens and workers are able to undergo diapause and overwinter; (4) the colony thus passes the winter without brood. This strategy, however, is not the most common.
The forms of dormancy which were found in ants extend from elementary quiescence to deep diapause. In exogenous-heterodynamic species, the diapause is optional for larvae and queens. The diapause occurs as a result of a direct reaction to temperature decline in the autumn but at a moment when the temperatures still exceed the developmental threshold.
On the contrary in endogenous-heterodynamic ant species, the diapause is compulsory for the colony and occurs eventually under any conditions. Two main factors restrict and control the internal brood-rearing cycle in these species. They are the endogenous timer and environmental conditions, temperature and photoperiod (in some species). But environmental cues can only regulate in some degree the moment of the onset of diapause by accelerating or delaying this event. All cold climate ants have adult diapause, so that their queens and workers are capable for overwintering. Queens and some workers experience diapause several times in their life. On the contrary, the ability of larvae to undergo diapause is not universal in ants. This is a major factor in seasonal cycle evolution in these insects.
The diapause of larvae in ants is facultative in most case. Temperature can affect larval development and induce diapause both directly and through the nurse workers. The larvae appeared to be entirely insensitive to the direct influence of photoperiods. The forms of social impact on larval diapause by workers are diverse in ants and range from nearly absolute control when the physiological state of workers completely defines the fate of larvae, to rather weak effects when in experimental conditions diapausing workers are unable to prevent the development of most overwintered larvae, and spring workers are capable to induce pupation of only a few diapausing autumn larvae.
We can conclude that the similar seasonal adaptations could arise in ant evolution independently many times and usually are not tightly bound to the taxonomic position of species. Nevertheless, several examples of certain seasonal cycle traits clearly confined to specific ant taxa have been found.
Acknowledgments
I am very grateful to students and members of the Department of Entomology, St. Petersburg State University, who participated in these long-term investigations in different years. This work was supported by grants from European Union INTAS program (94-2072), Russian Foundation of Basic Research (97-04-48987, 00-04-49003, 03-04-48854, 06-04-49383), Federal Program “Universities of Russia” (07.01.026, 07.01.327) and the Council for Grants from the President of the Russian Federation and for State Support of Leading Scientific Schools (00-15-97934, 2234.2003.4, 7130.2006.4).
\n',keywords:"Formicidae, ants, seasonal development, life cycles, homodynamic, heterodynamic, diapause, exogenous, endogenous, temperature, climate, evolution",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/60505.pdf",chapterXML:"https://mts.intechopen.com/source/xml/60505.xml",downloadPdfUrl:"/chapter/pdf-download/60505",previewPdfUrl:"/chapter/pdf-preview/60505",totalDownloads:1434,totalViews:222,totalCrossrefCites:1,totalDimensionsCites:2,totalAltmetricsMentions:0,impactScore:1,impactScorePercentile:61,impactScoreQuartile:3,hasAltmetrics:0,dateSubmitted:"June 20th 2017",dateReviewed:"February 13th 2018",datePrePublished:null,datePublished:"October 24th 2018",dateFinished:"April 4th 2018",readingETA:"0",abstract:"This chapter is a review of data on structure, diversity and adaptive properties of seasonal cycles in ants. Most tropical ants demonstrate homodynamic development. They do not show any developmental delays and all-year round the ontogenetic stages from egg to pupa exist in their nests. Some of the quasi-heterodynamic species have permeated into the regions with warm temperate climate but a true diapause did not evolve. Most temperate and all boreal climate ants are true heterodynamic. They manifest a real winter diapause (prospective dormancy) in their annual cycle. Thus, a variety of forms of dormancy, which were found in ants, extend from elementary quiescence to deep diapause. Heterodynamic ants use two main seasonal strategies with respect to brood rearing: strategy of concentrated brood rearing (Formica type) and strategy of prolonged brood rearing (Aphaenogaster type, Myrmica type). The larval stages at which diapause can occur are extremely variable among ants. The evolution of seasonal life cycles and possible ways of origin of diapause in ants are discussed. The subtropical (quasi-heterodynamic) and tropical (preadaptational) evolutionary paths to true heterodynamic development are considered. It is concluded that similar seasonal adaptations could arise in the evolution of ants independently many times and usually are not tightly bound to the taxonomic position of species.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/60505",risUrl:"/chapter/ris/60505",book:{id:"6400",slug:"the-complex-world-of-ants"},signatures:"Elena B. Lopatina",authors:[{id:"214873",title:"Associate Prof.",name:"Elena",middleName:null,surname:"Lopatina",fullName:"Elena Lopatina",slug:"elena-lopatina",email:"elena.lopatina@gmail.com",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",institution:{name:"Saint Petersburg State University",institutionURL:null,country:{name:"Russia"}}}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Peculiarities of seasonal development in ants as social insects",level:"1"},{id:"sec_3",title:"3. Homodynamic type of seasonal development",level:"1"},{id:"sec_4",title:"4. Heterodynamic cycles of development",level:"1"},{id:"sec_4_2",title:"4.1. Quasi-heterodynamic development",level:"2"},{id:"sec_5_2",title:"4.2. True heterodynamic development",level:"2"},{id:"sec_7",title:"5. Two seasonal strategies of brood rearing in heterodynamic ants",level:"1"},{id:"sec_7_2",title:"5.1. The strategy of prolonged brood rearing",level:"2"},{id:"sec_7_3",title:"5.1.1. More complete use of a favorable period for the development and available thermal resources",level:"3"},{id:"sec_8_3",title:"5.1.2. The ability to adapt to the duration of the warm period of a year by changing the amount of the rapid brood",level:"3"},{id:"sec_9_3",title:"5.1.3. The ability to stretch the development of larvae for two or even three summer seasons",level:"3"},{id:"sec_11_2",title:"5.2. The strategy of concentrated brood rearing (Formica type)",level:"2"},{id:"sec_13",title:"6. Review of heterodynamic annual cycles in ants",level:"1"},{id:"sec_13_2",title:"6.1. Seasonal cycles of Aphaenogaster type",level:"2"},{id:"sec_14_2",title:"6.2. Seasonal cycles of Myrmica type",level:"2"},{id:"sec_14_3",title:"6.2.1. Diapause in early (1st–3rd of 5 or 6) instars (Lepisiota, Plagiolepis, Tapinoma and some Camponotus)",level:"3"},{id:"sec_15_3",title:"6.2.2. Diapause in middle (2nd–4th of 5–6) instars (Camponotus s. str.)",level:"3"},{id:"sec_16_3",title:"6.2.3. Diapause in the late (3rd–4th of 4) instars (Harpagoxenus, Leptothorax, Temnothorax and Messor)",level:"3"},{id:"sec_17_3",title:"6.2.4. Diapause in the last (usually 3rd) instar (Manica, Diplorhoptrum, Leptanilla, Monomorium, Myrmica, Tetramorium)",level:"3"},{id:"sec_18_3",title:"6.2.5. Diapause in all (of 3–6) instars (sometimes except the first instar) (Aphaenogaster, Crematogaster, Lasius, Paratrechina, Camponotus, Tanaemyrmex)",level:"3"},{id:"sec_20_2",title:"6.3. Seasonal cycles of the Formica type",level:"2"},{id:"sec_22",title:"7. Two types of the regulation of heterodynamic annual cycles",level:"1"},{id:"sec_23",title:"8. The regulation of diapause in larvae",level:"1"},{id:"sec_24",title:"9. Evolution of annual cycles of development in ants",level:"1"},{id:"sec_24_2",title:"9.1. Subtropical (quasi-heterodynamic) path of evolution",level:"2"},{id:"sec_25_2",title:"9.2. Tropical (preadaptational) path of evolution",level:"2"},{id:"sec_26_2",title:"9.3. Taxonomic position and seasonal adaptations",level:"2"},{id:"sec_28",title:"10. Conclusions",level:"1"},{id:"sec_29",title:"Acknowledgments",level:"1"}],chapterReferences:[{id:"B1",body:'Brian M. Social Insects. Ecology and Behavioural Biology. London and New York: Chapman and Hall; 1983. 377 p. 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In: 12th Congress of the International Union for the Study of Social Insects, Paris, Sorbonne, August 21-27, 1994. 1994. p. 317'},{id:"B175",body:'Tauber M, Tauber C, Masaki S. Seasonal Adaptations of Insects. New York, Oxford: Oxford University Press; 1986. XV+411 p'},{id:"B176",body:'Danks H. Insect Dormancy: An Ecological Perspective. Ottawa: Biological Survey of Canada (Terrestrial Arthropods); 1987. IX+439 p'},{id:"B177",body:'Danks H. Key themes in the study of seasonal adaptations in insects II. Life-cycle patterns. Applied Entomology and Zoology. 2006;41:1-13. DOI: 10.1303/aez.2006.1'},{id:"B178",body:'Elmes G. The social biology of Myrmica ants. Actes des Colloques Insectes Sociaux. 1991;7:17-34'},{id:"B179",body:'Elmes G, Thomas J. Die Biologie und Ökologie der Ameisen Gattung Myrmica. In: Geiger W, editor. Tagfalter und ihre Lebensraume: Arten. Gefahrdung, Schutz, Basle: Schweizerischer Bund für Naturschutz; 1987. pp. 404-409'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Elena B. Lopatina",address:"elena.lopatina@gmail.com",affiliation:'
Department of Entomology, Saint Petersburg State University, Russian Federation
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1. Introduction
The truth is that since the arrival of Jan Van Reibeeck and his people in the Cape to date, South Africa’s land issue has never been as it was and it continued to be a big challenge for the majority of black people. From then onwards, the white people’s government was ensuring that the black people are alienated from all forms of owning land, as a way of subjecting and putting them under racial control. It was very clear that if the blacks were allowed to have land, their dependency on the western ways of life would be minimized, which would be a problem for colonization and its intended inequalities to perpetually take away the dignity of the black person. This is the correct argument of Resane [1] when contemplating being stripped of the land when a majority of people were removed and dumped into the designated areas demolished their life meaning, their dignity, civil rights, and respect. The agenda of transformation after many years of racial divisions and the subjection of the black majority in South Africa is one of the biggest but most challenging projects that many people can imagine. It is without any doubt that one of the most debated or discussed issues in South Africa in 2018 is land redistribution. It makes sense to also indicate that the transformation agenda in democratic South Africa for the past twenty-five years had been very slow in this regard; reconciliation of the racially and tribally divided nation is ultimately affected by this as part of the transformation. Moosa has the following to say in this regard: “While the issue of land can be used opportunistically by the politicians and their supporters, it is disingenuous to suggest that South Africans are unconcerned. The relationship of the land issue to reconciliation and inequality is visible for everyone to see.” [2].
This is evidenced by the racial and tribal attitudes that continue to haunt the South African society in churches, government departments, and workplaces. There is enough evidence to allude that the land redistribution program had been hit by delays, inaction, and contested political interest, while the ordinary citizens’ interests are being marginalised ([3], p. 1). It should be understood that the issue of whether the expropriation should be compensated or will not be discussed in this paper, but will argue that the land issue is a hindrance between South Africa and reconciliation.
It is therefore the purpose to argue how the land issue continues to marginalise the already marginalised people and open the gap of inequality even wider. It will be a waste of time if I pretend to encourage the political debates and different views which are embedded in the issue of land redistribution in this article; the discussion will generally focus on indicating how the delays in the land issue are a hindrance to the reconciliation that almost all people of South Africa would like to see between the whites and blacks. Despite the fact that the TRC headed by Desmond Tutu targeted a few people, it also did not do much for the reconciliation of the racially and tribally divided nation. My previous paper was on racism and tribalism and I tried to unveil how tribalism remains a thorn in the flesh of reconciliation. This paper also addresses racism which is raising its ugly head in the workplace, churches, and public square [4, 5].
2. Problem statement
We cannot easily identify our problems relating to any attempt of reconciliation project if we ignore to openly clarify that though the rifts of divisions existed in tribal lines, the focus is on the racial divisions by which apartheid managed very well to install successful laws that ensured that blacks were inferior towards the superior whites in many forms. The factors that are delaying the reconciliation of South African reconciliation for almost three decades are very complex, but for the sake of this specific study, the author intended to identify the slow pace at which the land redistribution is being administered as one issue. In discussing that, it will be noted that the difference that exists between the social groups existing in South Africa are in varying degrees, but the promise of the present democratic government even in the freedom charter gave the impression of equating all people that live in the land and promised this as one of the fundamental issues to play a role in the reconciliation process, for instance, Äll people shall govern” has no restriction for a particular social group within the borders of the country ([6], p. 1). The gist of reconciliation in this country must be seen from that context, particularly for this article.
The president of the republic, Mr Cyril Ramaphosa said:
“We cannot be a reconciled nation for as long as the majority of the people of South Africa continue to suffer from the injustices of the past. Access to land is a fundamental right of citizenship. It just not just empower communities and workers, it enhances food security, especially for people in the rural areas. Despite a comprehensive land reform programme, we have not made significant progress in this issue. Most of this country’s land remains in the hands of a few people in our country.” [7].
The sentiments of the president above resonate very well with his later statement in his first address as a state president when the ANC adopted the policy of land expropriation without compensation in its 54th Conference as he said:
We will accelerate our land redistribution programme not only to redress a grave historical injustice, but also to bring more producers into the agricultural sector and to make more land available for cultivation … this approach will include the expropriation of land without compensation. We are determined that expropriation without compensation should be implemented in a way that increases agricultural production, improves food security and ensure[s] that the land is returned to those from whom it was taken under colonialism and apartheid.5 [Emphasis added] (1–2) ([2], pp. 1–2).
These presidential statements and views show the clear direction of the ruling party, which is in line with the Freedom Charter, but the implantation is a different story that is met with contestations from different political parties. That on its own makes South African Reconciliation Barometer a public opinion that no one will be sure when it will come to practice.
According to SAPA ([8], p. 1), Trevor Manuel reported: “South Africa has almost the same number of people living in informal settlements now as it did in 1994. This view resonates with Ramantswana’s [9] articulation when saying:
The landlessness of the black masses is evidenced by the continuous mushrooming of informal settlements… To be landless in South African cities is to live in canals, under bridges, under trees, in parks, and in open spaces in front of buildings or shops. Even worse, to be a landless female is to be prey to slave-traders and sex-traders and to be first in line for endless rape. Landlessness sets up our people for dehumanization and exploitation.
This is even though the government provided three million houses during the period.” The minister went on to argue that in reversing this trend, it must also be known that South African cities were designed for the colonial elite; and it is a challenge to maintain that elitist live style with the challenges that are currently facing South Africa. Malema states as follows:
“The time for reconciliation is over. Now is the time for justice,” Malema told the parliament. “We must ensure that we restore the dignity of our people without compensating the criminals who stole our land.” (Business [10]).
The three statements from politicians above inform me of one main thing: we have a problem to reconcile the divided South Africans who have previously been divided and are still divided with the land distribution issue as one of the biggest stumbling blocks towards this goal. In other words, the reconciliation of this country will remain a dream that will never come true if the land issue is not dealt with properly . It is therefore the gist of this study to research and argue that land redistribution is essential for the stuck reconciliation programme which is part of the transformation agenda to carry on. If I have to use an understandable metaphor I would say that the bus is stuck. One of its wheels is termed “reconciliation; and this wheel is out because one of its bolts, “land redistribution,” has come out. Therefore, to get the bus moving means that we start with the lost bolts, by fixing them then the wheel (reconciliation) will be fitted to the bus and the bus will then start moving on. It does not matter how small or big the bolts are, they can stop the whole bus from moving. The reality is that South Africa is stuck in its transformation agenda, particularly in reconciliation and it demands theological knowhow, among others to try to make us going forward. The resistance and delays we are experiencing with the land issue undermine the broken black people. If reconciliation and justice go hand in hand, there is a great risk that South Africa will not achieve any reconciliation if the justice on a land issue is not dealt with to the satisfaction of the previously disadvantaged people. Let us face the fact that the landless people have lost human dignity, whose lives have lost meaning ([11], p. 3). Not only for pride’s sake, but without owning a land for African is a biggest humiliation one can imagine. For the author, it is like rubbing salt on the wound to speak about reconciliation between those who have land and those who do not have it. It is important to note that this paper did not intend to discuss politics, but to highlight the issue of land as a stumbling block, although unfortunately, the land issue has already been politicised in the country. This can be checked with the comments just above, which are made predominately by politicians.
3. Relevance of practical theology
Meiring [12] confirms that the problem is current and relevant when he says: “We do have to face the fact that South Africa, 20 years after democracy, is still a fractured and a very divided country. Racism, alienation, xenophobia are still with us, as is the case with corruption, greed and endemic violence.” This basic truth is what according to the author of this article dragged most of the transformational projects very slow, hence, for 25 years it looks like most of the citizen’s lives did not move for an inch from what it was before 1994.
It is important to note the fundamental truth in what Hall [13] warns us against when saying: “When we debate racism that it is important that we don’t only look at racism in relation to land as something that just happened in the past, but as an issue that continues to be felt today.” This argument informs us that as much as theology had tried to fight racism in the past, we need to remain alert that the fight is not over because the fight can only be won if all inhabitants of South African have an equal share in all opportunities, including land ownership. While theology is also caught ball watching, the patience of the people it intends to serve is also tried with by this resistance. The organisation called Landless People’s Movement (LPM) has come to alive because of the delay on the issue of land, as they are demanding answers to the very same issue. For me, inequality and any form of sabotage of human dignity is a concern that theology cannot keep silent about. Turning a blind eye on the situation is an indication of the demise of such a theology. One of the fathers of black theology, James Cone, spent much of his time on writing about this issue, when saying that any theology that does not answer the problems of the people at hand must cease to exist (Cone’s theology). According to Bosch ([14], 32–34), Jesus’ message on the kingdom includes God’s power that attacks the evils of the society in whatever form they manifest. The church of Jesus did not neglect works of compassion as part of proclaiming God’s message in totality; hence, pain, disease, demon and other forms of brokenness were dealt with. In addition, the intention of this paper is not to undermine the South African judiciary system and its procedures on issues of land and reconciliation, but to highlight and to unveil that the slow pace art in which the process is taking place is not helpful for many landless people. It is the very same democratic government that made the promise to deliver and it is the same government that must be held to account for their promises. For the government to do things legally and according to the rule of law is what everyone expects, but that does not necessarily imply that things must be as slow as they are with regards to the land issue. It is important for the author to mention that it is the very democratic government that must be held an account on the land issue because it is the one that made a promise to distribute the land at least by 2014 which is seven years back [15, 16]. Part of this article’s task is to critically highlight that it is the very same promise that must be kept, of course within the confines of the law that is governing the country. The author is not supportive of anything that is done outside the parameters of the law, urges that the required speed is maintained to distribute the land before things get out of hand as signs are already showing by some land grapping experienced.
4. Limitations and focus of this research
South Africa is one of the countries with young democracy after many years of colonialism and apartheid, implying that some of the changes taking place are still confusing and not perfectly done. The reality is that the challenges caused by inequality are complex and often intertwined, for instance, poverty, unemployment and other social challenges. It will be too ambitious to want to attend to all these in one chapter, but for the sake of this specific study, the author decided to avoid floating around all these problems, but focus on the land issue as a trap towards reconciliation. This does not however not mean that land issue will not affect other social problems or be affected by them, but the other social ills will also get their day in another research. This of course is not intended to claim that land issue will solve all the other existing problems, but it will open opportunities to face other realities, for instance, before one has a land it is difficult to imagine the exposure to issues like drought, storm and pest invasion. These are issues for future research.
Our African forefathers left us an idiom that says: “The best way to eat the elephant standing in your path is to cut it up into little pieces.” Slowly and patiently taking each problem at a time we will transform our country for the benefit of all who live in it.
5. Definition of some important concepts
The word land redistribution refers to a way in which the commercial land that has been owned by the whites is transferred to black South Africans. The literary translation of reconciliation suggests that it is a restoration of friendly relations; it can also be used as synonyms for appeasement, reunion, conciliation, harmonising and so forth. It is dangerous to use reconciliation a synonym of forgiveness because, according to Cornell [17], one can forgive, without providing an immediate reconciliation. Reconciliation is a restoration process whereby trust is deeply broken and it may be a lengthy process at times. The process of reconciliation depends on the attitude of the offender; the depth of betrayal and the pattern of offense. The unrepentant heart is difficult, if not impossible, to reconcile; hence, a line of difference should be drawn between restorative and retaliatory reconciliation.
The study made available by Akinyemi [15] on land ownership and usage for agriculture clearly makes a very good understanding of the link between landlessness and poverty. The author does not of course undermine the fact that there are many factors involved with regards to poverty, but since this is not the main intention of the study, it can be argued that having or not having the land can assist in making the separation between the rich and the poor. The author would avoid making generalisation if the division between the rich and poor for other countries also connects to land, but as for South Africa the research mentioned above makes that connection clear [15]. This however does dismiss the argument that every poverty in every country is alluded to land, that is not what this article is arguing or saying.
6. Background
To make the argument of this study more sounding, the Freedom Charter of 1995, from which all the promises were made by the liberation movements particularly with regards to land and governance will shed very important light for the cause of the frustrations that South Africans are having today. Three specific statements in the Freedom Charter are:
The Land Audit Report of November 2017 continued to emphasize by quoting the Constitution of the Republic by saying: “South Africa belongs to all who live in it, united in our diversity” [18]. The gist of this paper may be complicated for some readers without the knowledge of the Freedom Charter, which was a helping tool for the governing party and other liberal movements to gain the support of the majority in toppling the apartheid regime. It is on the background of these initial promises adopted in 1955 that the reactions of people on issues of land in South Africa can be attributed to.
There is a confirmation that the 2017 Land Audit Report stats on land reform were just a beginning of the long journey of redistributing the land by saying: “Finally, this exercise has revealed that we have just taken the first steps upon a long journey towards the goal of the sustainable relationship of South African citizens to one another through the effective management of land as a resource and nation-building”([19], p. 20).
According to [20], p. 1) the land restitution programme which is based on Section 25(7) of the Constitution,1 points that:
“A person or community dispossessed of property after 19 June 1913 as a result of past racially discriminatory laws or practices is entitled, to the extent provided by an Act of Parliament, either to restitution of that property or to equitable redress.” Some of the claims in this regard were attended to, but many are still outstanding and the landless are not aware of what the update is.
The state of affairs according to “The Borgen Project report” with regards to land occupation as in 2021 indicates that: “White South Africans accounted for less than 10% of the population after the apartheid in South Africa ended in 1994. However, 90% of white South Africans owned the land. In addition, about 72% of white South Africans owned farmland in 2017. Meanwhile, black South Africans owned only 4% of land and Indian South Africans owned about 5%. As such, poverty and land reform in South Africa remain large issues ([21], p. 1). This audit is supported by Phaliso [22] who exactly have the same statistics in his/her report dated 2018. The audit shows that whites owned the majority of land at 72%, followed by coloured people at 15%, Indians at 5% and Africans at 4%.
Until today, South Africa is still one of the countries with the highest levels of inequalities, according to World Bank 2019 statistics [23]. The disparities that exist between the rich and the poor is incomparable to most of the countries. From then onwards, the ownership of the land in the country was transferred into White hands, that continued even during the apartheid period [24]. The Natives Land Act which is regarded as apartheid’s original sin saw thousands of black families being forcefully removed from their land and since then onwards, black South African knew no peace with regards to those removals.
The continued removal of black people from the land went on even when homelands were formed in the late 1960s and became is a crack that has not yet mended even after close to three decades of democracy. The forceful removal of black ethnic groups aimed at giving some land for exclusive use had been and will always be blamed for the current inequalities. There is no doubt that the move to democracy in South Africa is challenged among others by the land reform process. This is because some effects of the apartheid regime were the historical unequal allocation of the land ([25], p. 1). These historical injustices should be dealt with for true reconciliation to surface. The truth which is undeniable, according to De Villiers [26] is that in four countries namely; Zimbabawe, Namibia, South Africa and Australia the historical land issues were characterised by the inequality, colonial dominance and discrimination which led to the indigenous inhabitants being forced off their land to the benefit of white settlers, even the ancestral lands where they buried their loved ones were forcefully taken from them. It is an undeniable fact that by whom and how the land is used has a very direct bearing on the growing unemployment as well as poverty statistics [27].
My background discussion starts with a quote by Butjwana Seokoma [28] who said:
“South Africa’s land reform programme, adopted by the African National Congress (ANC) led government in 1994, has a long way to go in redressing the historical injustice of land dispossession, denial of access to land and forced removals.”
It is evident that the problem about land redistribution in South Africa was caused by the attitude of the racial discrimination of the apartheid government, which most of us accept was inclusive of many policies that wronged our people to a great extent. This is when all tribal laws, tribal divisions and forceful removal of blacks from the areas designated for white use exclusively. The dispossession of land from black South Africans was a key part of the colonial and apartheid strategy of entrenching superiority over the black population, it rendered the blacks unable to live on their land, forcing them to work for the colonial and apartheid powers. It removed people from ancestral land, disabling them from being able to perform cultural rites, visit graves, bury the umbilical cords of newborns and the deceased on ancestral land. [29].
Sol Plaatjie, the first secretary of ANC, was quoted reiterating what the colonial ideology was all about saying: “Where will we get servants if Kaffirs (Africans) are allowed to become skilled? A kaffir with a thousand bags of wheat? If they are inclined to her the pedigree stock, let them improve their masters (colonial settlers) cattle and cultivate for the owner of the land, not for themselves.” (Motsoko [30]). Despite that the statement dealt a blow to the dignity and humanity of black people, this statement was manifested by ensuring that black people do not own land in this country; hence, the outcry today comes a long way.
For me the basic human right was violated just here and not much had been done to reverse it, particularly for ordinary people who lack even a small piece of land to erect a shack. On the other hand, “those who were favoured through legislation to acquire productive and at the expense of the disposed black majority were able to use that asset over a time as a leverage for wealth creation- to get ahead materially, to afford the comforts of life, to take their children to better schools, to build social resilience and to bequeath material legacy for future generations of their kin. The experience of the black majority was the complete opposite”(Daily Maverick, 14 August 2018).
It is the acknowledgement of all these wrongs that were supposed to be accompanied by a confession of guilt from those who intentionally benefited from this wrongdoing. From the inception of democratic government in 1994 this had been one of the slowest projects of transformation, since only a small percentage of land had been taken so far. The resistance and politicising of this aspect caused delays that see us loosing lives like in the apartheid regime, for instance people whose shacks are being washed away by floods due to lack of proper space to shelter themselves.
No one can deny that the dawn of the first democratic elections in 1994 was harnessed among others by the hope that the disadvantaged people of this country thought that it would change their lives, even in terms of having land to themselves. The honest response is that very little had been done and this, on its own, does not give courage that the ordinary citizens of this country will have land from the country of their birth. The dignity of the other is continuously delayed and denied.
Mr Petros Nkosi said:
“The land, our purpose is the land: that is what we must achieve. The land is our whole lives: we plough it for food, we build our houses from the soil, we live on it, and we are buried on it. When the whites took our land away from us, we lost the dignity of our lives, we could no longer feed our children, we were forced to become servants, we are treated like animals. Our people have many problems, we are beaten and killed by farmers, the wages we earn are too little to buy even a bag of mealie meal. We must unite together to help each other and face boers. But in everything we do we must remember that there is only one aim and one solution and that is the land, the soil, our world.” (18 May 2017), The constitutional court speaks about land and dignity). This is very similar to SAHRC’s position on land reform when they said: “Land is dignity and the restitution of land rights equals to restoration of dignity.”
This is the kind of the enmity and hatred that had been sowed by the issue of land, among others. We cannot continue blinding ourselves to think Mr Nkosi’s mind is his alone, since many black landless people are singing the same tune although they might not be very vocal. The dignity of a human being is on the land issue from birth to death. The previous state president Jacob Zuma also sang the same song of saying that it will be very difficult if not impossible to achieve true reconciliation until the land question is resolved ([2], p. 2).
If lack of land affects the dignity of the majority of this country, it means that by delaying and trying to avoid a quick movement towards it is also to deny that these people’s dignity must be returned to them. Justice delayed is justice denied. Therefore, how do we expect of the landowner to be reconciled to someone who does not even own a space where his/her shack is situated. It is the dignity of our people that is being tarnished by red ants who are evicting them from one place to the next on a daily basis. If my shack cannot give me enough privacy because there is no space between it and those of my neighbours , while the white farmer has enough space even for his dogs, cats and mice to play around, then the dignity of the one living in the shack cannot be equated to the dignity that the dogs and cats have. How is reconciliation possible? Parker ([31], p. 3) is very correct to mention that returning the land means restoring the dignity of the people. It should however be commended that one of the first steps towards land restitution began after the unbanning of liberation movements during the last days of the National Party’s rule in which FW De Klerk ‘s government abolished the Racially Based Land Measures Act 108 of 1991 where in a commission on land allocation was established ([26], p. 47). The situation as at 2019 according to [11] indicates that only 13% of South African is owned by black people while 87% of it is owned by whites.
We have two camps with regards to land issue in South Africa, those who have the land and the landless. Their walks of life, worldviews and life experiences still remain the same as it was more than twenty-five years ago; very opposed to each other and the question is: How do they join 16 December annual celebrations in the same fashion, while they remain in this situation? I am of the opinion that the reconciliation of people must not ignore the life world and levels of lives each person comes from. Some will people will argue that there are rich black people that own the houses that are on the first and second pictures, but I am fully aware that the majority of our people are still in the life indicated in the pictures below and yet they are not excluded from the discussions of reconciliation. To deal with and overcome the historical injustices of the past, where the land was seized under apartheid it is just imperative not to avoid addressing a land issue. That is what Gibson [24] in his book entitled Övercoming Historical injustices: Land Reconciliation in South Africa” is trying to argue. The land issue is undoubtedly at the heart of South African politics since it is an important cornerstone towards reconciling the nation that is haunted by racial divisions amongst others.
The first two pictures are of white farmers while the rest portray the living places for most black people living in South Africa.
The safety of the first two houses to natural disasters compared to the shacks in which one lives in fear, should high rainfalls come. This fear has been normalised to be a lived experience in which one is forced to create happiness despite circumstances.
Basic services: If one has to walk for kilometres to fetch “unhealthy water” meanwhile someone else’s swimming pool is always filled and the water may be healthier than the water the other is going to fetch.
Dwellings: The mountainous area is beautified by grass-covered roundavels which cannot even stand the test of some minor storms has become a great home in which most blacks spend four to five hours to reach and have their happiness there. An area that cannot have any resource at all, but life goes on as usual.
The filth and the sewerage smell cutting across shack area has been normalised into black life in the area, which can be a very serious concern for those living in the first two homes.
Space: The first two houses have enough space where even the dogs and cats can enjoy freedom while walking around, but that is categorically different from the experiences of the people living in the houses portrayed in third and fourth pictures.
In his unpublished paper entitled: “The power of Babel” Kritzinger spoke of the difference that has not disappeared between Jew and Greek, slave and free being language. The metaphor I would use is that of the language in the sense of the content of what people discuss. My opening question will be, What kind of language will be a talk between the people living in these two extremely different environments, for instance, the word water can mean something for the person whose swimming pool is always full of water meanwhile for the other person who is walking four to five kilometres just to fetch drinking water, a basic resource of life the same water may mean something different.
As if it is not enough, those find small space to just erect their shacks to live and are evicted by the red ants in the fifth picture, to add insult to injury.
The above pictures confirm what Siviwe Feketa mentioned in his article entitled: South Africa world’s most unequal society report’ when he reported:
The government has expressed disappointment with its track record of transforming the country after a World Bank report showed that inequality has deepened since the dawn of democracy, with the country being the most unequal society. The results of the probe, which assessed poverty and inequality from 1994 to 2015, revealed that only one in four South Africans could currently be stably considered as either middle class or upwards in terms of means. [32].
In Collins’ article entitled: “No vacant land in Joburg is safe from occupation,” he quoted one landless person in Alexander Township saying: “The empty stand represents an opportunity for dignity, privacy and a home she can finally call her own. It also represents a waste of something she has never had and is ready to fight for.” [33]. This is one example out of many or even the majority of the people in South Africa, whose dreams to have a land of their own in which they can build houses is a dream not coming true. It is difficult to imagine that for almost three decades after the democratic government which had one of its pillars in the freedom charter as to ensure that all people will access the land.
Many other issues can be raised by just comparing these pictures, but for reconciliation’s sake, there are many issues that would have to be attended to before we embark on the language of reconciling the two groups of people, unless the word reconciliation is only on our lips. Even though it is not the focus of this article, it is important to note that there are also squatter camps where white people are living, for instance, Munsieville in Krugersdorp. But it should however be known that this has been and is still a home for the majority of black South Africans. This has been part of the long history of inequality that was orchestrated by the elements of injustice and racism, among others. Recently, the very hot land debate was revived between the ruling party (ANC) and EFF where the bone of contention was that the government must be a custodian of the whole land while the ANC believed that they want to be custodians only of certain portions of the land in the country [34]. This of course have a close relationship with the argument of appropriation of land without compensation, which has been a headache for the government for a long time now. Of course, this very long debate is not of much interest in this study, but it is touched upon just to shed light as to where we are in terms of land ownership in South Africa, which has serious and should seriously continue to be one of the determining factors towards reconciling this very divided country.
7. Endangered lives continue to be normalised
I became concerned when even some black people who were supposed to have supported the brother in Mamelodi started criticising him and his family for his lost child when the pipes burst. I am referring to the case where Mike Mshiane and his Wife, Vinolia Sikele, lost a child when the water pipes burst below their five-year-old shack in Mamelodi East [35]. For me this incident took the child’s life because of the land problem more than anything else. My argument is that if the land redistribution programme had not been delayed and this couple got the land to put their shack or house on good land, their child would still be alive today. They were left without a choice, but to erect a shack in the wrong place because the land issue is still to take more decades to discuss. I do not want to talk about the Alexandra community, the shacks of whom are always washed away whenever floods come through that Jukskei river. The consistent warnings whenever heavy rains and storms are predicted are made to people living in low-lying areas like Alexander and others because it is known that their houses and shacks will be damaged and even taken away. McCain [36] confirms this in his report entitled “Heavy rain causes flooding in parts of Pretoria, with more rain expected.” In 2013 some homes of people were swept away by floods alongside the Tugela River in KZN because that was the only piece of land those people would call a home [37]. The lives of those without land remain at risk all the times. Without denying that these calamities and challenges happen in other countries also, the author’s argument is that within the context of landless people in South Africa, these would be avoided for some people if they had proper land to build their houses. Without evading the possibilities of natural calamities and disasters like drought, pests and invasions which the author is aware of, it is important to maintain the focus of this specific study, keeping in mind that the future research must attend to those issues for continuity’s sake. These are not the only challenges connecting to the topic, but poverty and unemployment as well, but for the sake of space and focus, the author will carry these into the future research.
It is the lives of the landless black people that are at risk there and yet they are expected to talk about reconciliation to someone who stays on a large portion of land. If we take a count of lives that were lost due to this delay of land redistribution, the statistics will show that the life of blacks is not a problem in South Africa. This level of inequality is also exposed during the lockdown due to the corona virus. Nocuze’s [38] picture where people were cuing a very long cure to the very small toilet of a certain Nolusindiso Xaka of Khayelitsha is a practical example. These people’s dignity is compromised while the lockdown measures of isolation are hindered. This cannot only be blamed on the lockdown measures, but also to the lack of land in which everyone could try to at least build his or her own toilet. It was also argued in that report that residents refused to move their shacks to make space for toilets, merely because there is no enough space left to do that ([38], p. 9).
8. Deprivation to basic services
The 06h00 am news on Wednesday 05 November 2019 from Munghana Lonene (SABC Tsonga Radio Station) included a painful story of people who are starting to dig their wells to get water in Hamanskraal, just about 30 minutes-drive northwards of Pretoria. This is an entirely black township. It should be noted that a large inequality across South African provinces regarding basic services such as piped water, toilets, formal dwelling signals how the structuring of this poverty was meant to benefit the white dwelling places. According to Gradin [39], this was structured so that it could also enhance the racial disparities and seriously deprive the black people from economic opportunities. It sounds like a joke to expect someone without a toilet or a bucket toilet to reconcile with someone who has more than five toilets in a house in which only two people live. Practically, there is no equal lifestyle there that can draw these two different families closer to one another. The dignity of the other is undermined from the onset. I will close this section by quoting Gradin : “In summary, the legacy of apartheid and colonisation has left Africans with several drawbacks that make them more likely to be poor, such as living in rural areas or in the poorest provinces, higher fertility, less education, and poorer labour market outcomes, even if it is difficult to determine which of them is more relevant than the others.” ([39], p. 194). In one of its broadcasting stations called “Munghana Lonene FM” the SABC indicated that in Merwe, a village outside Malamulele, people are sharing drinking water with their domestic animals (Munghana Lonene News, 06h30 on 11 October 2019). Another report from Limpopo indicated that a young girl, Humbelani Mudzanani was attacked by crocodiles and her body was found floating in the dam. This was because she was supposed to fetch water from the stream since they do not have water in the Tshitomboni village ([40], p. 9). The unfortunate and sad moment for this village is caused by the lack of delivery of one of the basic public services. It should be noted that while some are being killed by crocodiles for fetching water, others are not even fetching water since it is in their houses, and yet, we expect the two kinds of people to reconcile. It is difficult if not impossible to expect the person who sleeps in a linking shack with the one whose dogs are sleeping in a house with ceiling and air conditioner to be on the same level of live, let alone reconciliation. That is in line with what Feely, ([41], p. 1) argues when saying that dignity in every human being is intrinsic and cannot be separated from a human being. There is a clear indication that reconciliation which comes along with justice must not neglect the lost dignity of those who were stripped of the land by those who have the land. Although, the examples given to highlight the lack of service delivery may seem to be of a particular social or ethnic group, the author thinks that these are just examples, but the reality is that all ethnic groups are affected by this delivery. All nine provinces of South Africa have communities that are having such issues as water, sanitation and roads services. The most general one is that of the load shedding, which affects every citizen as well as visitors of the country. This can be summed up by Mazamisa’s ([42], p. 213) who called the situation “ a devastating effect on many communities and individuals.”
9. Fuelling of xenophobic attacks
I am convinced that whenever the land and resources are less than the consumers, the likelihood is a conflict in the form of xenophobia or afroforbia. There is one biblical story that can be used to explain this further. In Mark 6:35–40 the disciples of Jesus Christ had only five loaves of bread and two fishes, while the estimated crown on the day were about 500. Since it was humanly impossible for the available food to feed the large number of listeners, the disciples were quoted saying:
“This is a remote place,” they said, “and it’s already very late. Send the people away so that they can go to the surrounding countryside and villages and buy themselves something to eat.” This was a way to evade the responsibility of being a brother or a sister in terms of sharing the resources. (Mark 6, pp. 35–36).
The spirit of Ubuntu of sharing the little we have can often be challenged by the size of the resources we have. There are many ways of trying to evade responsibility when the challenge is that of sharing very limited resources. The current state of affairs in South Africa is characterised by the fact that it is those in the majority that do not have land. It does not mean that it is not a right for everyone to have a piece of land, but the government is in the same corner which the disciples of Jesus found themselves in when confronted by a large crowd with very little food. It is possible to think of solving the problem by sending the crowds home to enjoy the little resources without being faced by hungry people. Hatred among South Africans can also be fuelled by lack of land while others have it in abundance.
One of the reasons given for the xenophobic attacks in South Africa was that the foreigners are taking their jobs because they are ready to be paid peanuts. This is because the locals have trade unions that give the farm owners a tough time and instruct them to pay the workers decent living wages as well as giving them living conditions in their workplaces. Among others, many farm workers have been complaining that they stayed on the farms for many years and yet they do not have the right to own the land on which they can bury their loved ones. The experiences of the clash between residents and municipality law enforcement staff like those reported from Mfuleni has become a normal order of the day because the majority of black people do not have a piece of land just to put their houses or shacks. This is exactly what Merten [3] refers to in her report with sad pictures of the people with their belongings without roofs since they were buildings were destroyed, when she said: “Land ownership in South Africa remains a contentious issue, while calls for redistribution grow louder.” Marten also believes that an access to land is one of the keys to uplift unemployment. It is suspected of course, that one of the reasons the land distribution is a very slow issue it’s because the government loves money and they expect people to pay for the land, meanwhile the majority of the poor in the country are poor and unemployed. VeIi Hlatshwayo [43] was quoted saying: “The attacks against blacks by South Africans show that there is a need to continue the revolution. Until wealth and land is equally distributed among the citizens, such story will continue.” This is a clear indication that the reconciliation that is being expected in South Africa is still trapped in the web of the issue of land.
Besides xenophobic violence, the black local people often engage in battles over the land issue. In the programme entitled “Tiko axi etleri”the SABC Radio Station Munghana Lonene reported that one man was hospitalized as a result of the shooting that occurred between the two local village leadership over a land dispute outside Tzaneen. This is the village under the local chief Nkuna and the Mkgolobodo village which fought about a land grabbing until guns were used. The destruction of people’s homes and their goods was also mentioned (SABC, Tiko, 19 January 2022: 06h00-07h00).
10. Landlessness must be treated as a violation of the human right
The research by Naidoo and Naidoo [44] entitled “The erosion of human dignity in the New South Africa” explains amongst other things how humanity, particularly black humanity was unjustly robbed of their dignity and yet expected to embrace love. There is no way a land issue cannot be the top of the agenda of all sorts of injustices that put the black people in their adjacent poverty as they are today. Although some want to argue that we keep blaming the pre-1994 government whereas the country is now democratic, the author’s argument is that those who are still mourning their loss must be allowed to do so until such time when they realise that the reversal of the wrongs has been done. [45], p. 215) added to this argument by indicating that the very slow pace at which land reform is being done is cause for land grapping since many are losing patience to the process. For Goolam [46] the foundation of democracy, which is human dignity is attained by the equality of all people’s human rights and freedom. This clarifies why the argument that reconciliation is still an uphill to climb when other people’s dignity through the stripping of the land is still at stake.
11. Practical theological guidelines on land injustices
Theologically speaking, the God who created the land amongst other things is the same God of justice. Any theology that tries to evade responsibility by not listening to and addressing the cries of its immediate recipients is not only irrelevant, but must also cease to exist. The God of theology that we read about in the Bible is the creator of heavens and earth. He also created human beings in His own image before placing them in the garden, which is part of the land (Genesis 1, pp. 26–28). If God wanted people to be landless He would not have placed them on land and instructed them to subdue it (Gen.1:28). The creation was all perfect when the inclusion of human being placed on the land was part of the creation story. In other words, the statement that it was very good in Genesis 1:31 was vocalised after the placing of human being on land was included as part of the plan of God. The author’s interpretation is that the creation story is incomplete or imperfect without the man being given the land to subdue it. For that reason, it became a painful part of history when human beings were expelled from the allocated land after the disobedience of Genesis 3. In practical sense, this implies that it is the very same Creator who has powers to remove from any part of land whoever He placed, not another human being.
It is clear that some researchers pointed out that there was a serious role that religion, particularly John Calvin’s Reformed theology legitimized the policies of racial segregation that put all the ([47], p. 9). This on its own suggests that as much as the religious and faith communities have played a role in using their doctrines and other influence for this unjust practice to be perpetuated, they are not to be left out when reversing the very same issues they were supporting. The author of this article sees the importance of revising and unteaching that which was taught as an essential tool towards the different that will topple down the inhuman segregation, including on land issue. In that way the wrong role of the church will be reversed and it is not only about writing of confessing, but also by involving the church into these negotiations. The rebuilding of the community through restoration of human dignity is not only political move, but a moral and ethical obligation for which the church has a part to play [48]. For mobilisation and socialisation of the community faith community must play their part, for example through documents like that of Smith [49] which was circulated in the Dutch Reformed Church Synod around 2002. The document was entitled: Land Reform and the church ([47], p. 12). Such documents do not only influence the community, but they also can be given a chance for suggestions to the policy makers for consideration in rebuilding the nation as the advancement of the land redistribution is called upon. The influence of organisations like SACC is still respected amongst faith-based organisations and that is a positive point, they should use their influence to get involved and not stay back when these negotiations as well as land grabbing are taking place. Using the scriptures the task of advocacy and prophetic voice in challenging the state about this may make the difference. Making of submissions and petitions to portfolio committees and the relevant government ministries falls within the democratic right of every organisation and individuals, these opportunities have not been much exploited.
For ([11], p. 1) besides its political controversies, the land has also emotional and religious attachment to African people, hence theology cannot afford to be left out in the issue of land. It is for this reason that the author’s understanding links very well with that of Mlambo when indicating that the struggle for land is a struggle for justice because these two have been tied together ([50], p. 1). Theology is incomplete if it avoids including the issue of land as part of its discourse. The thoughts of the late Prof ([51], p. 2) when he aimed to look at the characteristics of land theology, which is one of the most important catalysts of black theology of liberation. There is no way the colonial justice of cultural domination and oppression that forcefully took the land can be justified. God has and will always been on the side of the poor and the oppressed, hence it is convincing that He cannot be the author of such injustice. If the unjust taking of the land from black people is one of the causes of oppression that demanded a revolutionary struggle, then the very logical argument is that there cannot be a reconciliation without liberation ([52], p. 6). This is very in line with Chimhanda [53] who has seen that black theologians have recognized the connection that exists between achieving authentic peace, liberation and reconciliation. These concepts plus justice cannot be separated in political terms, but they should move together for true reconciliation to be achieved. According to Takatso Mofokeng [54] in his chapter “Land is our mother” the symbolical mother is ready to save her child at all costs. The clear lesson we can learn is of the suffering of the child who is orphaned by the loss of the mother. This is the kind of toleration landless and poor South African had been subjected to for all these years. It only makes sense to understand the land as an integral part of African view of life [51].
There is a need for Liberation theology to play its role. The liberation of Israelites from Egypt was incomplete before the occupation of the promised land; hence, God continued to move with them through deserts into Canaan. This is why God was still actively involved with His chosen nation during the conquest of Canaanites inhabitants under the military leadership of Joshua to ensure that they indeed settle on the land. It is my understanding that the 1994 democratic elections were just phased one of victory, but phase two must still take place where the complete restoration of the land to the black people must happen. God knows not to rest until the land occupation is still not on the side of the black masses who are still landless in South Africa. One of the fathers of Liberation theology, James Cone [55] wrote much about the liberative message of God where he argues that God is on the side of the poor and the oppressed. In John 10:10 we read: “The thief comes only to steal and kill and destroy; I have come that they may have life, and have it to the full.” It is a contention of the author of this article that life is not in its fullness to those who do not have land, rights dignity and respect they deserve.
If there was a painful part in the history of Israel as a chosen nation of God, it was that period of exile. One of the main reasons why being exiled was a painful experience was that people would be removed from the land of their own into a foreign land. It has always been a prayer and desire for Israelites that would be returned into the land of their own, the land promised to their ancestor Abraham. It was for the reason his ancestral land and its gates were destroyed that Nehemiah (1:1–5) sat down and wept.
From a theological point of view there are clear biblical stories that tell that God the Creator dislike and punishes those who wrongly take the land in support of those who lost it. According to ref. [1](p. 184) the biblical story of how Ahab’s greedy over the vineyards of Naboth makes a lot of relevance here. While king Ahab was insisting that Naboth should sell him his vineyards he continuously indicated to him that it was against his tradition to give away his inheritance. Our theology must teach us that there is a close relationship between the land and what our forefathers left for us. The anger of God on those who forcefully take the land of others was demonstrated when God pronounced a judgement in I Kings 21:19, which was fulfilled in 1 Kings 22:37–38. This judgement can be justified by Deuteronomy 19:14 which says: “Do not move your neighbor’s boundary stone set up by your predecessors in the inheritance you receive in the land the LORD your God is giving you to possess.” True theology must teach and confront people of South Africa, including the government with this reality. According to James Cone, God is always on the side of the poor and oppressed, in this context, on the side of those who does not have land meanwhile their forefathers were citizens in the same country. Theologians and the church must be able to inform the government that it is both an abuse of power and against God’s will to keep landlessness a normal way of life ([56], p. 53).
There is amongst others, one biblical example of justice towards the issue of the land that was wrongfully taken from its owner. The story of the Shunamite woman whose land was unjustly taken was restored and her dignity was restored in 2 Kings 8:5–6 which says: And while he was telling the king how Elisha had restored the dead to life, behold, the woman whose son he had restored to life appealed to the king for her house and her land. And Gehazi said, “My lord, O king, here is the woman, and here is her son whom Elisha restored to life.” 6 And when the king asked the woman, she told him. So the king appointed an official for her, saying, “Restore all that was hers, together with all the produce of the fields from the day that she left the land until now.” This is in line with Cone’s theology when he says that God is always on the side of the poor and afflicted. The South African situation of the black masses on land issue is not exceptional. If the South African government is serious about reconciling South Africans, their dignity should be looked at also in the lenses of land issue as a matter of urgency. The current South African context demands that we have the kind of prophetic liberative theology like the one we read above. The government should take the lead in ensuring that before the land is grabbed like it was happening recently, justice is done on the issues of land redistribution.
Reconciliation demands that both parties should move from their own comfort zones and meet at the middle point. In other words, readiness of reaching compromise should be encouraged from both sides, those who have it and those who do not have it. ([4], p. 5) argues that reconciliation can be achieved by trying to give more than what one receives. White landowners must come out instead of remaining in the dark and let the politicians only fight their battle. Jacob and Esau met at a middle point for reconciliation to take place. The nature of reconciliation comes with the recognition and confession of the wrongs of the past to embrace a new future. The defence mechanisms that we are seeing in South Africa will never yield the reconciliation that we need to see. Baloyi further indicated that the power of faith, sacrifice and moving from comfort zone into the meeting point of the divided people would help towards reconciliation ([5], p. 6). It is important to note that as difficult as the situation is, there are some people and churches that give land to people as a way of bringing peace and this should be commended. These people are willing to lose what they have to at least play role in bringing peace.
Paul also touched on the issue of reconciliation in his teachings. According to [57], p. 1), Paul’s doctrine of reconciliation involves individual, corporate, cosmic and eschatological dimensions. According to Paul, reconciliation becomes the objective work of God through Jesus Christ in 2 Cor.5:19. In Colossians 1:19–21 it is written:
For God was pleased to have all his fullness dwell in him, and through him to reconcile to himself all things, whether things on earth or things in heaven, by making peace through his blood, shed on the cross. Once you were alienated from God and were enemies in your minds because off your evil behaviour. But now he has reconciled you by Christ’s physical body through death to present you holy in his sight, without blemish and free from accusation.
The word reconciliation finds its roots in the Pauline letters in the Bible. The hostilities that existed between the Jews and Greeks are also dealt with in a reconciliatory spirit from Galatians 3:28. He uses the basis of this reconciliation on the work done on the cross. Basically, the important argument he uses is that reconciliation and salvation should move alongside each other [58, 59, 60].
Although, it does not necessarily mean that the one will cause the other, I support Louw’s ([61], p. 9) opinion that forgiveness and reconciliation should go alongside each other. If the South African societies have not forgiven one another, just like it is being evidenced by racial attacks in churches, workplaces and public squares, it remains very difficult to conceptualise some form of reconciliation. It is just a pity that, according to Volf [62], the churches are presumed to be instruments of the peace we want to see become the enemies of that peace by influencing conflicts. Wielenga ([63], p. 7) suggest that the churches should plan to have story-sharing encounters around a meal or braai in the church regularly.
According to Stamps ([64], p. 1812), the death of Jesus on the cross to reconcile people with God must be a lesson to teach us that the church must stand up and fight for justice and reconciliation. Although forgiveness and reconciliation are not the same things, they should move along [62]. I must add that it becomes difficult, if not impossible, to expect reconciliation and forgiveness where justice did not prevail. To add on this Wielega ([63], p. 9) urges the churches to plan an open space where people can freely express and share their past stories as a healing method.
Lederack’s model of reconciliation- From the Algerian unequal context Lederach’s proposals can also be discussed in trying to shape how reconciliation in the divided South Africa can be mapped going forward. Without detailing much of what he said, it is important to make use of concepts like “mercy”, “peace” and “justice” must be allowed to play an important role in intensifying reconciliation and unity into a fragmented society like South Africa ([65], p. 30). The author of this paper suggests an inclusive approach when addressing the issue of land redistribution. This is because there are evidences that their processes of land are often frustrated by lack of proper consultation with all the interested parties, from both black and white people of this country. Kgatle [66] indicated in his research entitled…. that racial comments keep on being posted even in our social media, which play a role in frustrating the process since unfounded general labelling can cause resistance from those who can meaningfully contribute to the discussion and process. Although Kgatle [66] argued this in the context of the Faith Mission Church in South Africa, the bigger context of the country is also characterised by what he meant. Instead, the author would advocate for the spirit of accountability towards another human life. The Biblical message of being each other’s keepers apply also to this issue. Those with land, particularly unused land should start thinking about other human beings whose shacks and houses are planted along the river beds while others are cramped in a line of shacks without basic service because of being in a small space.
Pastoral leadership can also play a role by influencing the local traditional leaders to call upon people to strategize together towards land reforming instead of participating in unofficial land grabbing. Traditional leaders have an opportunity to raise these issues to the government officials who always come down to seek their influence for political elections.
12. Conclusion
The disparities and inequalities between the previously disadvantaged South Africans and those who were privileged by the apartheid regime are still a visible one today. Not much had been done to close the gap between those who had life and those who did not have it. Therefore, when starting to talk about reconciliation we need to be careful since the two classes of people are practically far from each other. The main item at the centre cannot be ignored, land redistribution which will enable the previously disadvantaged to start climbing the ladder. Ignorance to this issue is like sitting on the wheels of the transformation agenda of reconciliation. The reality is that besides it being a political agenda, there is a strong need for reconciliation to the “still” divided South African nation. It should be noted that reconciliation is part of the bigger transformation agenda which demands the reversal of “all” the injustices of the past. These injustices include the land issue which remains a thorn in the flesh of many South Africans. It is humanly difficult to imagine how the society with such a great inequality can be reconciled without serious consideration of basic issues like land redistribution, which is a serious need for the black majority.
\n',keywords:"reconciliation, black people, poverty, informal settlement, landlessness",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/81631.pdf",chapterXML:"https://mts.intechopen.com/source/xml/81631.xml",downloadPdfUrl:"/chapter/pdf-download/81631",previewPdfUrl:"/chapter/pdf-preview/81631",totalDownloads:6,totalViews:0,totalCrossrefCites:0,dateSubmitted:"February 14th 2022",dateReviewed:"March 7th 2022",datePrePublished:"May 9th 2022",datePublished:null,dateFinished:"May 4th 2022",readingETA:"0",abstract:"South Africa is a country with a history of racial divides and those divides are still visible today. One of the many issues that characterise such divisions is inequality with regard to land ownership. The bigger part of the land is still owned by the minority white people while the blacks, who are the majority; or previously disadvantaged people, are still landless. This is evident from the escalation of informal settlements in areas surrounding the cities of Johannesburg and Pretoria, which have millions of black people cramped into small areas in shacks while most whites are enjoying large portions of land with few people to live on it. Many black people even lack land for shelter. On the other hand, the message of reconciling the country is being made loud to both who has the land and those who do not have it. Therefore, it becomes a serious challenge to imagine and look for reconciling strategies while these two camps still have this unresolved issue. The dilemma that is faced with is how will the different camps embrace the same message while they remain in these different situations. It is argued in this article that justice on the land issue should be first attended to so that it serves as a door to reconciliation.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/81631",risUrl:"/chapter/ris/81631",signatures:"Baloyi Magezi Elijah",book:{id:"11429",type:"book",title:"Sustainability, Ecology, and Religions of the World",subtitle:null,fullTitle:"Sustainability, Ecology, and Religions of the World",slug:null,publishedDate:null,bookSignature:"Dr. Levente Hufnagel",coverURL:"https://cdn.intechopen.com/books/images_new/11429.jpg",licenceType:"CC BY 3.0",editedByType:null,isbn:"978-1-80355-436-5",printIsbn:"978-1-80355-435-8",pdfIsbn:"978-1-80355-437-2",isAvailableForWebshopOrdering:!0,editors:[{id:"10864",title:"Dr.",name:"Levente",middleName:null,surname:"Hufnagel",slug:"levente-hufnagel",fullName:"Levente Hufnagel"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}},authors:null,sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Problem statement",level:"1"},{id:"sec_3",title:"3. Relevance of practical theology",level:"1"},{id:"sec_4",title:"4. Limitations and focus of this research",level:"1"},{id:"sec_5",title:"5. Definition of some important concepts",level:"1"},{id:"sec_6",title:"6. Background",level:"1"},{id:"sec_7",title:"7. Endangered lives continue to be normalised",level:"1"},{id:"sec_8",title:"8. Deprivation to basic services",level:"1"},{id:"sec_9",title:"9. Fuelling of xenophobic attacks",level:"1"},{id:"sec_10",title:"10. Landlessness must be treated as a violation of the human right",level:"1"},{id:"sec_11",title:"11. Practical theological guidelines on land injustices",level:"1"},{id:"sec_12",title:"12. Conclusion",level:"1"}],chapterReferences:[{id:"B1",body:'Resane KT. Naboth’s vineyard: Theological lessons for the South African Land ISSUE. Acta Theologica. 2015;35(1):174-188'},{id:"B2",body:'Moosa M. 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Sustainable Reconciliation in Divided Societies. Washington DC: Unites States Institute of Peace; 1998'},{id:"B66",body:'Kgatle MS. A practical theological approach to the challenge of poverty in post-1994 South Africa: Apostolic Faith Mission as a case study. Herv Teology Studies. 2017;73(3):1-9'}],footnotes:[{id:"fn1",explanation:"This article is a reworked paper delivered in a M&D seminar at University of Pretoria on 28 January 2019."}],contributors:[{corresp:"yes",contributorFullName:"Baloyi Magezi Elijah",address:"baloye@unisa.ac.za",affiliation:'
Department of Philosophy, Practical and Systematic Theology, University of South Africa, South Africa
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The photo-excited current is charged in photodiode junction capacitance itself. The current is changed to the voltage about 1012 times without feedback resistance. The minimum detectable power of InGaAs CIA system with liquid nitrogen was achieved 0.1 fW of 10 sec integration time at the wavelength of 1.28 μm. The optical band pass filter-based system for ultra–low-level light detection was succeeded in spectrum measurement of 1O2 by 13-LOOH with cytochrome c. The 8 channel InGaAs-CIA array system enables to achieve optical multichannel detection for ultra-low level light at 10−13 W from 10−15 W level in the near-infrared region. The optical resolution was about 200 nm by 1 channel. The spectrum of 1O2 by mixing NaOCl and H2O2 was demonstrated. The shape of spectrum by 1O2 was matched to that of measured by the spectrometer. The system was succeeded in instantaneous 1O2 spectrum measurement without moving the wavelength dispersion device. The generation of 1O2 by photo-excited Rose Bengal was fabricated to develop food antioxidant chemistry or source reagent of cosmetic product. The system uses super luminosity LED for excitation light source and InGaAs CIA. The 1O2 generation will be controlled by the InGaAs-CIA monitoring system. 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Definition of Terms:
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Book - collection of Works distributed in a book format, whose selection, coordination, preparation, and arrangement has been performed and published by IntechOpen, and in which the Work is included in its entirety in an unmodified form along with one or more other contributions, each constituting separate and independent sections, but together assembled into a collective whole.
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Work - a book Chapter (as well as Conference Papers), including any and all content, graphics, images and/or other materials forming part of, or accompanying, the Chapter/Conference Paper.
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Attribution – appropriate credit for the used Work or book.
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Creative Commons licenses – enable licensors to retain copyright while allowing others to use their Works in an appropriate way.
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Rules of Attribution for Works Published by IntechOpen
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With the purpose of protecting Authors' copyright and the transparent reuse of OA (Open Access) content, IntechOpen has developed Rules of Attribution of Works licensed under Creative Commons licenses.
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All Chapters published in IntechOpen books prior to October 2011 are licensed under the Creative Commons Attribution-NonCommercial-ShareAlike 3.0 Unported license (CC BY-NC-SA 3.0);
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All Chapters published in IntechOpen books after October 2011 are licensed under the Creative Commons Attribution 3.0 Unported license (CC BY 3.0);
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In case you reuse or republish any of the Works licensed under CC licenses, you must abide by the guidelines outlined below:
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1. Rules for reusing of books in their entirety or significant parts of books
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All rights to Books and other compilations published on the IntechOpen platform and in print are reserved by IntechOpen. The Copyright to Books and other compilations is subject to a separate Copyright from any that exists in the included Works.
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A Book in its entirety or a significant part of a Book cannot be translated freely without specific written consent by the publisher. Further information can be obtained at permissions@intechopen.com.
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In instances where permission is obtained from the publisher for reusing or republishing the Book, or significant parts of the Book, all of the following conditions apply:
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Information about the first publisher must be provided – please note the fact that the material was originally published by IntechOpen as an OA (Open Access) publication must be acknowledged;
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All original Academic Editor(s) must be credited;
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Since you are reusing content that someone else created and allowed you to use freely, you must credit all Authors involved;
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The type of license that is available for the Works must be indicated, as well as a link to the license provided, so that others can investigate the terms of the license. You will be aware that the material can be used for free in consequence of the CC license attribution, so you must acknowledge that fact. It is not sufficient that the material is Creative Commons, because that says nothing about how the material can actually be used. There are different CC licenses and you have to identify the specific license that is being used;
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Any original Copyright Notices associated, with the Works which constitute the Book must be kept intact;
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Provision of the original title of the Book, as well as the original titles of any individual Works;
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Provision of the URL where the Book is hosted, with a notice to the effect that the Book is an OA (Open Access) publication;
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Provision of the URL to every individual Work which constitutes the Book with a notice that the Work is an OA (Open Access) publication. As the material has been accessed for free, it is incumbent upon you to provide the source so that others can also access it for free.
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\\n\\n
Every single Work that is used has to be attributed in the way described. If you are unsure about proper attribution, please write to permissions@intechopen.com.
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2. Rules of attribution for works published by IntechOpen
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Individual Works originally published in IntechOpen books are licensed under Creative Commons licenses and can be freely used under terms of the respective CC license, if properly attributed. In order to properly attribute the Work you must respect all the conditions outlined below:
\\n\\n
\\n\\t
Credit all Authors – since you are reusing contents that someone created and allowed you to use freely, you have to acknowledge authorship;
\\n\\t
Indicate the type of license under which the Work is available and provide the URL to the license so others can find out the license terms. Preferably keep intact any original Copyright Notice associated with the Chapter (if any). You will be aware that the material can be used for free in consequence of the CC license attribution, so you must acknowledge that fact. It is not sufficient that the material is Creative Commons, because that says nothing about how the material can actually be used. There are different CC licenses and you have to identify the specific license that is being used;
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Provide the URL where the Work is hosted, preferably providing the original title of the Work, as well as the original title of the Book with a notification that the Work is an OA (Open Access) publication. As the material has been accessed for free, it is incumbent upon you to provide the source so that others can also access it for free;
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Provide information about the first publisher – please note the fact that the material was originally published by IntechOpen as an OA (Open Access) Work must be acknowledged.
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Every single Work that is used has to be attributed in the way as described. If you are unsure about proper attribution, please contact Us at permissions@intechopen.com.
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In the event that you use more than one of IntechOpen's Works published in one or more books (but not a significant part of the book that is under separate Copyright), each of these have to be properly attributed in the way described.
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IntechOpen does not have any claims on newly created copyrighted Works, but the Works originally published by IntechOpen must be properly attributed.
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All these rules apply to BOTH online and offline use.
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Parts of the Rules of Attribution are based on Work Attributing Creative Commons Materials published by the Australian Research Council Centre of Excellence for Creative Industries and Innovation, in partnership with Creative Commons Australia, which can be found at creativecommons.org.au licensed under Creative Commons Attribution 2.5 Australia license, and Best practices for attribution published by Creative Commons, which can be found at wiki.creativecommons.org under the Creative Commons Attribution 4.0 license.
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All the above rules are subject to change, IntechOpen reserves the right to take appropriate action if any of the conditions outlined above are not met.
Work - a book Chapter (as well as Conference Papers), including any and all content, graphics, images and/or other materials forming part of, or accompanying, the Chapter/Conference Paper.
\n\n
Attribution – appropriate credit for the used Work or book.
\n\n
Creative Commons licenses – enable licensors to retain copyright while allowing others to use their Works in an appropriate way.
\n\n
Rules of Attribution for Works Published by IntechOpen
\n\n
With the purpose of protecting Authors' copyright and the transparent reuse of OA (Open Access) content, IntechOpen has developed Rules of Attribution of Works licensed under Creative Commons licenses.
\n\n
\n\t
All Chapters published in IntechOpen books prior to October 2011 are licensed under the Creative Commons Attribution-NonCommercial-ShareAlike 3.0 Unported license (CC BY-NC-SA 3.0);
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All Chapters published in IntechOpen books after October 2011 are licensed under the Creative Commons Attribution 3.0 Unported license (CC BY 3.0);
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\n\n
In case you reuse or republish any of the Works licensed under CC licenses, you must abide by the guidelines outlined below:
\n\n
1. Rules for reusing of books in their entirety or significant parts of books
\n\n
All rights to Books and other compilations published on the IntechOpen platform and in print are reserved by IntechOpen. The Copyright to Books and other compilations is subject to a separate Copyright from any that exists in the included Works.
\n\n
A Book in its entirety or a significant part of a Book cannot be translated freely without specific written consent by the publisher. Further information can be obtained at permissions@intechopen.com.
\n\n
In instances where permission is obtained from the publisher for reusing or republishing the Book, or significant parts of the Book, all of the following conditions apply:
\n\n
\n\t
Information about the first publisher must be provided – please note the fact that the material was originally published by IntechOpen as an OA (Open Access) publication must be acknowledged;
\n\t
All original Academic Editor(s) must be credited;
\n\t
Since you are reusing content that someone else created and allowed you to use freely, you must credit all Authors involved;
\n\t
The type of license that is available for the Works must be indicated, as well as a link to the license provided, so that others can investigate the terms of the license. You will be aware that the material can be used for free in consequence of the CC license attribution, so you must acknowledge that fact. It is not sufficient that the material is Creative Commons, because that says nothing about how the material can actually be used. There are different CC licenses and you have to identify the specific license that is being used;
\n\t
Any original Copyright Notices associated, with the Works which constitute the Book must be kept intact;
\n\t
Provision of the original title of the Book, as well as the original titles of any individual Works;
\n\t
Provision of the URL where the Book is hosted, with a notice to the effect that the Book is an OA (Open Access) publication;
\n\t
Provision of the URL to every individual Work which constitutes the Book with a notice that the Work is an OA (Open Access) publication. As the material has been accessed for free, it is incumbent upon you to provide the source so that others can also access it for free.
\n
\n\n
Every single Work that is used has to be attributed in the way described. If you are unsure about proper attribution, please write to permissions@intechopen.com.
\n\n
2. Rules of attribution for works published by IntechOpen
\n\n
Individual Works originally published in IntechOpen books are licensed under Creative Commons licenses and can be freely used under terms of the respective CC license, if properly attributed. In order to properly attribute the Work you must respect all the conditions outlined below:
\n\n
\n\t
Credit all Authors – since you are reusing contents that someone created and allowed you to use freely, you have to acknowledge authorship;
\n\t
Indicate the type of license under which the Work is available and provide the URL to the license so others can find out the license terms. Preferably keep intact any original Copyright Notice associated with the Chapter (if any). You will be aware that the material can be used for free in consequence of the CC license attribution, so you must acknowledge that fact. It is not sufficient that the material is Creative Commons, because that says nothing about how the material can actually be used. There are different CC licenses and you have to identify the specific license that is being used;
\n\t
Provide the URL where the Work is hosted, preferably providing the original title of the Work, as well as the original title of the Book with a notification that the Work is an OA (Open Access) publication. As the material has been accessed for free, it is incumbent upon you to provide the source so that others can also access it for free;
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Provide information about the first publisher – please note the fact that the material was originally published by IntechOpen as an OA (Open Access) Work must be acknowledged.
\n
\n\n
Every single Work that is used has to be attributed in the way as described. If you are unsure about proper attribution, please contact Us at permissions@intechopen.com.
\n\n
In the event that you use more than one of IntechOpen's Works published in one or more books (but not a significant part of the book that is under separate Copyright), each of these have to be properly attributed in the way described.
\n\n
IntechOpen does not have any claims on newly created copyrighted Works, but the Works originally published by IntechOpen must be properly attributed.
\n\n
All these rules apply to BOTH online and offline use.
\n\n
Parts of the Rules of Attribution are based on Work Attributing Creative Commons Materials published by the Australian Research Council Centre of Excellence for Creative Industries and Innovation, in partnership with Creative Commons Australia, which can be found at creativecommons.org.au licensed under Creative Commons Attribution 2.5 Australia license, and Best practices for attribution published by Creative Commons, which can be found at wiki.creativecommons.org under the Creative Commons Attribution 4.0 license.
\n\n
All the above rules are subject to change, IntechOpen reserves the right to take appropriate action if any of the conditions outlined above are not met.
\n\n
Policy last updated: 2016-06-09
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Kim"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"7661",title:"Heat and Mass Transfer",subtitle:"Advances in Science and Technology Applications",isOpenForSubmission:!1,hash:"c29b5c2ce24925a935ca52b8344fbb99",slug:"heat-and-mass-transfer-advances-in-science-and-technology-applications",bookSignature:"Alfredo Iranzo",coverURL:"https://cdn.intechopen.com/books/images_new/7661.jpg",editedByType:"Edited by",editors:[{id:"67352",title:"Dr.",name:"Alfredo",middleName:null,surname:"Iranzo",slug:"alfredo-iranzo",fullName:"Alfredo Iranzo"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}],booksByTopicTotal:2,seriesByTopicCollection:[],seriesByTopicTotal:0,mostCitedChapters:[{id:"67726",doi:"10.5772/intechopen.86322",title:"CFD Simulation of Heat and Mass Transfer for Climate Control in Greenhouses",slug:"cfd-simulation-of-heat-and-mass-transfer-for-climate-control-in-greenhouses",totalDownloads:1120,totalCrossrefCites:3,totalDimensionsCites:3,abstract:"Greenhouse plant production involves a number of processes such as transpiration, condensation, photosynthesis, and climate control. Such processes, in turn, set off mass and heat transfer phenomena that influence not only the quality and quantity of crop production but also its environmental cost. While these processes have considerably been analyzed in separate, they strongly interact with one another. For instance, increased radiation (mainly thermal infrared) increases temperature, reduces humidity, consequently increases transpiration, and affects CO2 exchange as well as other reaction rates. Computational fluid dynamics (CFD) is a numerical tool with a solid physical basis which allows, through the construction of a computational model, to simulate the fluid flow environment. Heating, ventilation, and condensation have been analyzed in the greenhouse environment with CFD techniques. The current challenge is the interaction of these processes and their impact on the production system. The present work summarizes some CFD investigations carried out in this topic, in order to analyze the processes of heat and mass transfer in a greenhouse for agronomic purposes.",book:{id:"7661",slug:"heat-and-mass-transfer-advances-in-science-and-technology-applications",title:"Heat and Mass Transfer",fullTitle:"Heat and Mass Transfer - Advances in Science and Technology Applications"},signatures:"Cruz Ernesto Aguilar Rodriguez and Jorge Flores Velazquez",authors:[{id:"173578",title:"Dr.",name:"Jorge",middleName:null,surname:"Flores-Velazquez",slug:"jorge-flores-velazquez",fullName:"Jorge Flores-Velazquez"}]},{id:"66158",doi:"10.5772/intechopen.84706",title:"Numerical Solution to Two-Dimensional Freezing and Subsequent Defrosting of Logs",slug:"numerical-solution-to-two-dimensional-freezing-and-subsequent-defrosting-of-logs",totalDownloads:620,totalCrossrefCites:3,totalDimensionsCites:3,abstract:"Two-dimensional mutually connected mathematical models have been created, solved, and verified for the transient non-linear heat conduction in logs during their freezing and subsequent defrosting. The models reflect the influence of the internal sources of latent heat of both the free and bound water on the logs’ freezing process and also the impact of the temperature on the fiber saturation point of wood species, with whose participation the current values of the thermo-physical characteristics in each separate volume point of the subjected to freezing and subsequent defrosting logs are computed. The chapter presents solutions of the models with explicit form of the finite-difference method and their validation towards own experimental studies. Results from experimental and simulative investigation of 2D non-stationary temperature distribution in the longitudinal section of beech and pine logs with a diameter of 0.24 m and length of 0.48 m during their many hours freezing in a freezer and subsequent defrosting at room temperature are presented, visualized, and analyzed.",book:{id:"7661",slug:"heat-and-mass-transfer-advances-in-science-and-technology-applications",title:"Heat and Mass Transfer",fullTitle:"Heat and Mass Transfer - Advances in Science and Technology Applications"},signatures:"Nencho Deliiski and Natalia Tumbarkova",authors:[{id:"43040",title:"Prof.",name:"Nencho",middleName:"Stanev",surname:"Deliiski",slug:"nencho-deliiski",fullName:"Nencho Deliiski"},{id:"284649",title:"Dr.",name:"Natalia",middleName:"Yordanova",surname:"Tumbarkova",slug:"natalia-tumbarkova",fullName:"Natalia Tumbarkova"}]},{id:"67626",doi:"10.5772/intechopen.86738",title:"The Boundary Element Method for Fluctuating Active Colloids",slug:"the-boundary-element-method-for-fluctuating-active-colloids",totalDownloads:920,totalCrossrefCites:0,totalDimensionsCites:2,abstract:"The boundary element method (BEM) is a computational method particularly suited to solution of linear partial differential equations (PDEs), including the Laplace and Stokes equations, in complex geometries. The PDEs are formulated as boundary integral equations over bounding surfaces, which can be discretized for numerical solution. This manuscript reviews application of the BEM for simulation of the dynamics of “active” colloids that can self-propel through liquid solution. We introduce basic concepts and model equations for both catalytically active colloids and the “squirmer” model of a ciliated biological microswimmer. We review the foundations of the BEM for both the Laplace and Stokes equations, including the application to confined geometries, and the extension of the method to include thermal fluctuations of the colloid. Finally, we discuss recent and potential applications to research problems concerning active colloids. The aim of this review is to facilitate development and adoption of boundary element models that capture the interplay of deterministic and stochastic effects in the dynamics of active colloids.",book:{id:"8416",slug:"non-equilibrium-particle-dynamics",title:"Non-Equilibrium Particle Dynamics",fullTitle:"Non-Equilibrium Particle Dynamics"},signatures:"William E. Uspal",authors:[{id:"279308",title:"Prof.",name:"William",middleName:null,surname:"Uspal",slug:"william-uspal",fullName:"William Uspal"}]},{id:"66487",doi:"10.5772/intechopen.85735",title:"Mean Aspects Controlling Supercritical CO2 Precipitation Processes",slug:"mean-aspects-controlling-supercritical-co-sub-2-sub-precipitation-processes",totalDownloads:736,totalCrossrefCites:1,totalDimensionsCites:2,abstract:"The use of supercritical CO2 is an excellent alternative in extraction, particle precipitation, impregnation and reaction processes due to its special properties. Solubility of the compound in supercritical CO2 drives the precipitation process in different ways. In supercritical antisolvent process, mass and heat transfers, phase equilibria, nucleation, and growth of the compound to be precipitated are the main phenomena that should be taken into account. Mass transfer conditions the morphology and particle size of the final product. This transfer could be tuned altering operating conditions. Heat transfer in non-isothermal process influences on mixing step the size of generated microparticles. In rapid expansion of supercritical solution, phenomena as the phase change from supercritical to a CO2 gas flow, rapid mass transfer and crystallization of the compound, and expansion jet define the morphology and size of the final product. These phenomena a priori could be modulated tuning a large number of operating parameters through the experiments, but the correlations and modeling of these processes are necessary to clarify the relative importance of each one. Moreover, particle agglomeration in the expansion jet and CO2 condensation are determinant phenomena which should be avoided in order to conserve fine particles in the final product.",book:{id:"7661",slug:"heat-and-mass-transfer-advances-in-science-and-technology-applications",title:"Heat and Mass Transfer",fullTitle:"Heat and Mass Transfer - Advances in Science and Technology Applications"},signatures:"Antonio Montes, Clara Pereyra and Enrique J. Martínez de la Ossa",authors:[{id:"55991",title:"Mr.",name:"Antonio",middleName:null,surname:"Montes",slug:"antonio-montes",fullName:"Antonio Montes"},{id:"55992",title:"Dr.",name:"Clara",middleName:null,surname:"Pereyra",slug:"clara-pereyra",fullName:"Clara Pereyra"},{id:"55993",title:"Dr.",name:"Enrique",middleName:null,surname:"Martinez De La Ossa",slug:"enrique-martinez-de-la-ossa",fullName:"Enrique Martinez De La Ossa"}]},{id:"66317",doi:"10.5772/intechopen.85254",title:"Review Heat Transfer of Non-Newtonian Fluids in Agitated Tanks",slug:"review-heat-transfer-of-non-newtonian-fluids-in-agitated-tanks",totalDownloads:1001,totalCrossrefCites:1,totalDimensionsCites:1,abstract:"The heating and cooling of non-Newtonian liquids in tanks with mechanical impellers are operations commonly employed as chemical reactors, heat exchangers, distillers, extractors, thinners and decanters. In particular, the design of heat exchangers (jackets, helical coils, spiral coils and vertical tubular baffles) in tanks requires the prior knowledge of the rheology of the liquid for the calculation of the convection coefficients and the Reynolds number, in order to obtain the area thermal exchange. This chapter aimed to present the basic concepts of tanks with agitation, non-Newtonian liquids, hydrodynamics, heat transfer and, finally, with a practical design example for engineers and undergraduate students.",book:{id:"7661",slug:"heat-and-mass-transfer-advances-in-science-and-technology-applications",title:"Heat and Mass Transfer",fullTitle:"Heat and Mass Transfer - Advances in Science and Technology Applications"},signatures:"Vitor da Silva Rosa and Deovaldo de Moraes Júnior",authors:[{id:"187128",title:"Ph.D.",name:"Vitor",middleName:null,surname:"Rosa",slug:"vitor-rosa",fullName:"Vitor Rosa"},{id:"188792",title:"Dr.",name:"Deovaldo",middleName:null,surname:"Moraes Júnior",slug:"deovaldo-moraes-junior",fullName:"Deovaldo Moraes Júnior"}]}],mostDownloadedChaptersLast30Days:[{id:"66878",title:"Design of Industrial Falling Film Evaporators",slug:"design-of-industrial-falling-film-evaporators",totalDownloads:1753,totalCrossrefCites:0,totalDimensionsCites:0,abstract:"The high performance evaporators are important for process industries such as food, desalination and refineries. The falling film evaporators have many advantages over flooded and vertical tubes that make them best candidate for processes industries application. The heat transfer area is the key parameter in designing of an evaporator and many correlations are available to estimate the size of tube bundle. Unfortunately, most of the correlation is available only for pure water and above 322 K saturation temperatures. Out of these conditions, the areas are designed by the extrapolation of existing correlations. We demonstrated that the actual heat transfer values are 2–3-fold higher at lower temperature and hence simple extrapolated estimation leads to inefficient and high capital cost design. We proposed an accurate heat transfer correlation for falling film evaporators that can capture both, low temperature evaporation and salt concentration effectively. It is also embedded with unique bubble-assisted evaporation parameter that can be only observed at low temperature and it enhances the heat transfer. The proposed correlation is applicable from 280 to 305 K saturation temperatures and feed water concentration ranges from 35,000 to 95,000 ppm. The uncertainty of measured data is less than 5% and RMS of regressed data is 3.5%. In this chapter, first part summarized the all available correlations and their limitations. In second part, falling film evaporation heat transfer coefficient (FFHTC) is proposed and model is developed. In the last part, experimentation is conducted and FFHTC developed and compared with conventional correlations.",book:{id:"7661",slug:"heat-and-mass-transfer-advances-in-science-and-technology-applications",title:"Heat and Mass Transfer",fullTitle:"Heat and Mass Transfer - Advances in Science and Technology Applications"},signatures:"Muhammad Wakil Shahzad, Muhammad Burhan and Kim Choon Ng",authors:[{id:"174208",title:"Dr.",name:"Muhammad Wakil",middleName:null,surname:"Shahzad",slug:"muhammad-wakil-shahzad",fullName:"Muhammad Wakil Shahzad"},{id:"249811",title:"Dr.",name:"Muhammad",middleName:null,surname:"Burhan",slug:"muhammad-burhan",fullName:"Muhammad Burhan"},{id:"254696",title:"Prof.",name:"Kim Choon",middleName:null,surname:"Ng",slug:"kim-choon-ng",fullName:"Kim Choon Ng"}]},{id:"66102",title:"Heat and Mass Transfer of Additive Manufacturing Processes for Metals",slug:"heat-and-mass-transfer-of-additive-manufacturing-processes-for-metals",totalDownloads:1302,totalCrossrefCites:0,totalDimensionsCites:0,abstract:"Additive manufacturing (AM), a method in which a part is fabricated layer by layer from a digital design package, provides the potential to produce complex components at reduced cost and time. Many techniques (using many different names) have been developed to accomplish this via melting or solid-state joining. However, to date, only a handful can be used to produce metallic parts that fulfill the requirements of industrial applications. The thermal physics and weld pool behaviors in metal AM process have decisive influence on the deposition quality, the microstructure and service performance of the depositions. Accurate analysis and calculation of thermal processes and weld pool behaviors are of great significance to the metallurgy analysis, stress and deformation analysis, process control and process optimization etc. Numerical modeling is also a necessary way to turn welding from qualitative description and experience-based art into quantitative analysis- and science-based engineering branch. In this chapter, two techniques for producing metal parts are explored, with a focus on the thermal science of metal AM: fluid flow and heat transfer. Selective laser melting (SLM) is the one that is most widely used because it typically has the best resolution. Another is named metal fused-coated additive manufacturing (MFCAM) that is cost competitive and efficient in producing large and middle-complex components in aerospace applications.",book:{id:"7661",slug:"heat-and-mass-transfer-advances-in-science-and-technology-applications",title:"Heat and Mass Transfer",fullTitle:"Heat and Mass Transfer - Advances in Science and Technology Applications"},signatures:"Zhengying Wei and Jun Du",authors:[{id:"47614",title:"Prof.",name:"Zhengying",middleName:null,surname:"Wei",slug:"zhengying-wei",fullName:"Zhengying Wei"},{id:"282052",title:"Dr.",name:"Jun",middleName:null,surname:"Du",slug:"jun-du",fullName:"Jun Du"}]},{id:"66563",title:"Heat and Mass Transfer in Outward Convex Corrugated Tube Heat Exchangers",slug:"heat-and-mass-transfer-in-outward-convex-corrugated-tube-heat-exchangers",totalDownloads:1037,totalCrossrefCites:0,totalDimensionsCites:1,abstract:"Heat and mass transfer in outward convex corrugated tube heat exchangers is of significant importance for the optimization, fabrication, and application of outward convex corrugated tube heat exchangers. This chapter gives a deep investigation of the heat and mass transfer in outward convex corrugated tube heat exchangers. Based on the experimental setup developed, the performances of a novel outward convex corrugated tube heat exchanger are presented. Simulation methods are then used to detail the heat and mass transfer at tube side and shell side of the outward convex corrugated tube heat exchanger, and these include the flow structure, temperature distribution, and turbulence kinetic energy. Heat and mass transfer enhancements of the outward convex corrugated tube heat exchanger are also studied, and they are from tube side, shell side, and overall system aspects. Finally, multi-objective optimization of the outward convex corrugated tube heat exchanger is conducted to obtain the optimal performances through using Response Surface Methodology (RSM) and Non-dominated Sorting Genetic Algorithm (NSGA-II). Main conclusions and future outlook are then briefly stated and summarized. We firmly believe that the contents presented in this chapter can not only enrich the knowledge of heat exchangers but also develop methods for studying heat exchangers.",book:{id:"7661",slug:"heat-and-mass-transfer-advances-in-science-and-technology-applications",title:"Heat and Mass Transfer",fullTitle:"Heat and Mass Transfer - Advances in Science and Technology Applications"},signatures:"Huaizhi Han, Bingxi Li, Yaning Zhang, Quan Zhu and Ruitian Yu",authors:[{id:"23828",title:"Dr.",name:"Quan",middleName:null,surname:"Zhu",slug:"quan-zhu",fullName:"Quan Zhu"},{id:"148369",title:"Prof.",name:"Bingxi",middleName:null,surname:"Li",slug:"bingxi-li",fullName:"Bingxi Li"},{id:"196928",title:"Dr.",name:"Yaning",middleName:null,surname:"Zhang",slug:"yaning-zhang",fullName:"Yaning Zhang"},{id:"281875",title:"Prof.",name:"Huaizhi",middleName:null,surname:"Han",slug:"huaizhi-han",fullName:"Huaizhi Han"},{id:"282268",title:"Mr.",name:"Ruitian",middleName:null,surname:"Yu",slug:"ruitian-yu",fullName:"Ruitian Yu"}]},{id:"66317",title:"Review Heat Transfer of Non-Newtonian Fluids in Agitated Tanks",slug:"review-heat-transfer-of-non-newtonian-fluids-in-agitated-tanks",totalDownloads:1001,totalCrossrefCites:1,totalDimensionsCites:1,abstract:"The heating and cooling of non-Newtonian liquids in tanks with mechanical impellers are operations commonly employed as chemical reactors, heat exchangers, distillers, extractors, thinners and decanters. In particular, the design of heat exchangers (jackets, helical coils, spiral coils and vertical tubular baffles) in tanks requires the prior knowledge of the rheology of the liquid for the calculation of the convection coefficients and the Reynolds number, in order to obtain the area thermal exchange. This chapter aimed to present the basic concepts of tanks with agitation, non-Newtonian liquids, hydrodynamics, heat transfer and, finally, with a practical design example for engineers and undergraduate students.",book:{id:"7661",slug:"heat-and-mass-transfer-advances-in-science-and-technology-applications",title:"Heat and Mass Transfer",fullTitle:"Heat and Mass Transfer - Advances in Science and Technology Applications"},signatures:"Vitor da Silva Rosa and Deovaldo de Moraes Júnior",authors:[{id:"187128",title:"Ph.D.",name:"Vitor",middleName:null,surname:"Rosa",slug:"vitor-rosa",fullName:"Vitor Rosa"},{id:"188792",title:"Dr.",name:"Deovaldo",middleName:null,surname:"Moraes Júnior",slug:"deovaldo-moraes-junior",fullName:"Deovaldo Moraes Júnior"}]},{id:"65692",title:"Advances in Concentrated Solar Power: A Perspective of Heat Transfer",slug:"advances-in-concentrated-solar-power-a-perspective-of-heat-transfer",totalDownloads:1114,totalCrossrefCites:0,totalDimensionsCites:0,abstract:"Solar energy has the potential to reduce the dependence on the dwindling supply of fossil fuels through concentrated solar power (CSP) technology. CSP plants utilize solar thermal energy to produce electrical energy based on different thermodynamic power cycles. Solar collectors, reflectors, receivers, thermal fluid, and turbines are the main components of each CSP plant and involve intensive heat transfer at all stages. This chapter illustrates the thermal characteristics of the main components used in CSP technology. In addition, the solar thermal fluid characteristics and its stable operational ranges are discussed in this chapter. Heat capacity, vapor pressure, volume expansion, density and viscosity of the thermal fluid should not differ significantly at different temperatures during various operation stages because these variations can cause failure in the system, which is designed at the fixed material properties. Currently, CSP technology is associated with a higher cost compared to the electricity generated through gas power plants. Many efforts are made to search for sustainable and inexpensive materials to minimize the cost of CSP. One critical issue faced by CSP technology is the intermittent nature of the sun. Modern CSP plants integrate thermal energy storage (TES) unit to smoothen the power production or to shift the production from peak sunshine hours to peak demand hours.",book:{id:"7661",slug:"heat-and-mass-transfer-advances-in-science-and-technology-applications",title:"Heat and Mass Transfer",fullTitle:"Heat and Mass Transfer - Advances in Science and Technology Applications"},signatures:"Fadi Alnaimat and Yasir Rashid",authors:[{id:"151722",title:"Dr.",name:"Fadi",middleName:null,surname:"Alnaimat",slug:"fadi-alnaimat",fullName:"Fadi Alnaimat"},{id:"291252",title:"Mr.",name:"Yasir",middleName:null,surname:"Rashid",slug:"yasir-rashid",fullName:"Yasir Rashid"}]}],onlineFirstChaptersFilter:{topicId:"954",limit:6,offset:0},onlineFirstChaptersCollection:[],onlineFirstChaptersTotal:0},preDownload:{success:null,errors:{}},subscriptionForm:{success:null,errors:{}},aboutIntechopen:{},privacyPolicy:{},peerReviewing:{},howOpenAccessPublishingWithIntechopenWorks:{},sponsorshipBooks:{sponsorshipBooks:[],offset:8,limit:8,total:0},allSeries:{pteSeriesList:[{id:"14",title:"Artificial Intelligence",numberOfPublishedBooks:8,numberOfPublishedChapters:87,numberOfOpenTopics:6,numberOfUpcomingTopics:0,issn:"2633-1403",doi:"10.5772/intechopen.79920",isOpenForSubmission:!0},{id:"7",title:"Biomedical Engineering",numberOfPublishedBooks:12,numberOfPublishedChapters:98,numberOfOpenTopics:3,numberOfUpcomingTopics:0,issn:"2631-5343",doi:"10.5772/intechopen.71985",isOpenForSubmission:!0}],lsSeriesList:[{id:"11",title:"Biochemistry",numberOfPublishedBooks:27,numberOfPublishedChapters:286,numberOfOpenTopics:4,numberOfUpcomingTopics:0,issn:"2632-0983",doi:"10.5772/intechopen.72877",isOpenForSubmission:!0},{id:"25",title:"Environmental Sciences",numberOfPublishedBooks:1,numberOfPublishedChapters:9,numberOfOpenTopics:4,numberOfUpcomingTopics:0,issn:"2754-6713",doi:"10.5772/intechopen.100362",isOpenForSubmission:!0},{id:"10",title:"Physiology",numberOfPublishedBooks:11,numberOfPublishedChapters:139,numberOfOpenTopics:4,numberOfUpcomingTopics:0,issn:"2631-8261",doi:"10.5772/intechopen.72796",isOpenForSubmission:!0}],hsSeriesList:[{id:"3",title:"Dentistry",numberOfPublishedBooks:8,numberOfPublishedChapters:129,numberOfOpenTopics:0,numberOfUpcomingTopics:2,issn:"2631-6218",doi:"10.5772/intechopen.71199",isOpenForSubmission:!1},{id:"6",title:"Infectious Diseases",numberOfPublishedBooks:13,numberOfPublishedChapters:106,numberOfOpenTopics:3,numberOfUpcomingTopics:1,issn:"2631-6188",doi:"10.5772/intechopen.71852",isOpenForSubmission:!0},{id:"13",title:"Veterinary Medicine and Science",numberOfPublishedBooks:9,numberOfPublishedChapters:101,numberOfOpenTopics:3,numberOfUpcomingTopics:0,issn:"2632-0517",doi:"10.5772/intechopen.73681",isOpenForSubmission:!0}],sshSeriesList:[{id:"22",title:"Business, Management and Economics",numberOfPublishedBooks:1,numberOfPublishedChapters:11,numberOfOpenTopics:2,numberOfUpcomingTopics:1,issn:null,doi:"10.5772/intechopen.100359",isOpenForSubmission:!0},{id:"23",title:"Education and Human Development",numberOfPublishedBooks:0,numberOfPublishedChapters:0,numberOfOpenTopics:2,numberOfUpcomingTopics:0,issn:null,doi:"10.5772/intechopen.100360",isOpenForSubmission:!1},{id:"24",title:"Sustainable Development",numberOfPublishedBooks:0,numberOfPublishedChapters:9,numberOfOpenTopics:4,numberOfUpcomingTopics:1,issn:null,doi:"10.5772/intechopen.100361",isOpenForSubmission:!0}],testimonialsList:[{id:"13",text:"The collaboration with and support of the technical staff of IntechOpen is fantastic. The whole process of submitting an article and editing of the submitted article goes extremely smooth and fast, the number of reads and downloads of chapters is high, and the contributions are also frequently cited.",author:{id:"55578",name:"Antonio",surname:"Jurado-Navas",institutionString:null,profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002bRisIQAS/Profile_Picture_1626166543950",slug:"antonio-jurado-navas",institution:{id:"720",name:"University of Malaga",country:{id:null,name:"Spain"}}}},{id:"6",text:"It is great to work with the IntechOpen to produce a worthwhile collection of research that also becomes a great educational resource and guide for future research endeavors.",author:{id:"259298",name:"Edward",surname:"Narayan",institutionString:null,profilePictureURL:"https://mts.intechopen.com/storage/users/259298/images/system/259298.jpeg",slug:"edward-narayan",institution:{id:"3",name:"University of Queensland",country:{id:null,name:"Australia"}}}}]},series:{item:{id:"24",title:"Sustainable Development",doi:"10.5772/intechopen.100361",issn:null,scope:"
\r\n\tTransforming our World: the 2030 Agenda for Sustainable Development endorsed by United Nations and 193 Member States, came into effect on Jan 1, 2016, to guide decision making and actions to the year 2030 and beyond. Central to this Agenda are 17 Goals, 169 associated targets and over 230 indicators that are reviewed annually. The vision envisaged in the implementation of the SDGs is centered on the five Ps: People, Planet, Prosperity, Peace and Partnership. This call for renewed focused efforts ensure we have a safe and healthy planet for current and future generations.
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\r\n\tThis Series focuses on covering research and applied research involving the five Ps through the following topics:
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\r\n\t1. Sustainable Economy and Fair Society that relates to SDG 1 on No Poverty, SDG 2 on Zero Hunger, SDG 8 on Decent Work and Economic Growth, SDG 10 on Reduced Inequalities, SDG 12 on Responsible Consumption and Production, and SDG 17 Partnership for the Goals
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\r\n\t2. Health and Wellbeing focusing on SDG 3 on Good Health and Wellbeing and SDG 6 on Clean Water and Sanitation
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\r\n\t3. Inclusivity and Social Equality involving SDG 4 on Quality Education, SDG 5 on Gender Equality, and SDG 16 on Peace, Justice and Strong Institutions
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\r\n\t4. Climate Change and Environmental Sustainability comprising SDG 13 on Climate Action, SDG 14 on Life Below Water, and SDG 15 on Life on Land
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\r\n\t5. Urban Planning and Environmental Management embracing SDG 7 on Affordable Clean Energy, SDG 9 on Industry, Innovation and Infrastructure, and SDG 11 on Sustainable Cities and Communities.
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\r\n\tThe series also seeks to support the use of cross cutting SDGs, as many of the goals listed above, targets and indicators are all interconnected to impact our lives and the decisions we make on a daily basis, making them impossible to tie to a single topic.
",coverUrl:"https://cdn.intechopen.com/series/covers/24.jpg",latestPublicationDate:"April 24th, 2022",hasOnlineFirst:!0,numberOfPublishedBooks:0,editor:{id:"262440",title:"Prof.",name:"Usha",middleName:null,surname:"Iyer-Raniga",slug:"usha-iyer-raniga",fullName:"Usha Iyer-Raniga",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002bRYSXQA4/Profile_Picture_2022-02-28T13:55:36.jpeg",biography:"Usha Iyer-Raniga is a professor in the School of Property and Construction Management at RMIT University. Usha co-leads the One Planet Network’s Sustainable Buildings and Construction Programme (SBC), a United Nations 10 Year Framework of Programmes on Sustainable Consumption and Production (UN 10FYP SCP) aligned with Sustainable Development Goal 12. The work also directly impacts SDG 11 on Sustainable Cities and Communities. She completed her undergraduate degree as an architect before obtaining her Masters degree from Canada and her Doctorate in Australia. Usha has been a keynote speaker as well as an invited speaker at national and international conferences, seminars and workshops. Her teaching experience includes teaching in Asian countries. She has advised Austrade, APEC, national, state and local governments. She serves as a reviewer and a member of the scientific committee for national and international refereed journals and refereed conferences. She is on the editorial board for refereed journals and has worked on Special Issues. Usha has served and continues to serve on the Boards of several not-for-profit organisations and she has also served as panel judge for a number of awards including the Premiers Sustainability Award in Victoria and the International Green Gown Awards. Usha has published over 100 publications, including research and consulting reports. Her publications cover a wide range of scientific and technical research publications that include edited books, book chapters, refereed journals, refereed conference papers and reports for local, state and federal government clients. She has also produced podcasts for various organisations and participated in media interviews. She has received state, national and international funding worth over USD $25 million. Usha has been awarded the Quarterly Franklin Membership by London Journals Press (UK). Her biography has been included in the Marquis Who's Who in the World® 2018, 2016 (33rd Edition), along with approximately 55,000 of the most accomplished men and women from around the world, including luminaries as U.N. Secretary-General Ban Ki-moon. 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Other positions she has held at the university include Vice-Dean of Master Programs, Vice-Dean of the Degree in Biology and Vice-Dean for Mobility and Enterprise and Engagement at the Faculty of Science (University of Alicante). She received her Bachelor in Biology in 1998 (University of Alicante) and her PhD in 2003 (Biochemistry, University of Alicante). She undertook post-doctoral research at the University of East Anglia (Norwich, U.K. 2004-2005; 2007-2008).\nHer multidisciplinary research focuses on investigating archaea and their potential applications in biotechnology. She has an H-index of 21. She has authored one patent and has published more than 70 indexed papers and around 60 book chapters.\nShe has contributed to more than 150 national and international meetings during the last 15 years. Her research interests include archaea metabolism, enzymes purification and characterization, gene regulation, carotenoids and bioplastics production, antioxidant\ncompounds, waste water treatments, and brines bioremediation.\nRosa María’s other roles include editorial board member for several journals related\nto biochemistry, reviewer for more than 60 journals (biochemistry, molecular biology, biotechnology, chemistry and microbiology) and president of several organizing committees in international meetings related to the N-cycle or respiratory processes.",institutionString:null,institution:{name:"University of Alicante",institutionURL:null,country:{name:"Spain"}}},editorTwo:null,editorThree:null},{id:"15",title:"Chemical Biology",coverUrl:"https://cdn.intechopen.com/series_topics/covers/15.jpg",isOpenForSubmission:!0,editor:{id:"441442",title:"Dr.",name:"Şükrü",middleName:null,surname:"Beydemir",slug:"sukru-beydemir",fullName:"Şükrü Beydemir",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0033Y00003GsUoIQAV/Profile_Picture_1634557147521",biography:"Dr. Şükrü Beydemir obtained a BSc in Chemistry in 1995 from Yüzüncü Yıl University, MSc in Biochemistry in 1998, and PhD in Biochemistry in 2002 from Atatürk University, Turkey. 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He is a member of the Turkish Biochemical Society, American Chemical Society, and German Genetics society. Dr. Ekinci published around ninety scientific papers, reviews and book chapters, and presented several conferences to scientists. He has received numerous publication awards from several scientific councils. 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Since 1983, he has been a faculty member of the RO Perelman Department of Dermatology, NYU School of Medicine, where he is codirector of a training grant in cutaneous biology. Dr. Blumenberg’s research is focused on the epidermis, expression of keratin genes, transcription profiling, keratinocyte differentiation, inflammatory diseases and cancers, and most recently the effects of the microbiome on the skin. 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The applications of this research cover many related fields, such as biotechnology and medicine, where, for example, Bioinformatics contributes to faster drug design, DNA analysis in forensics, and DNA sequence analysis in the field of personalized medicine. Personalized medicine is a type of medical care in which treatment is customized individually for each patient. Personalized medicine enables more effective therapy, reduces the costs of therapy and clinical trials, and also minimizes the risk of side effects. Nevertheless, advances in personalized medicine would not have been possible without bioinformatics, which can analyze the human genome and other vast amounts of biomedical data, especially in genetics. The rapid growth of information technology enabled the development of new tools to decode human genomes, large-scale studies of genetic variations and medical informatics. 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