Barely three months into the new year and we are happy to announce a monumental milestone reached - 150 million downloads.
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This achievement solidifies IntechOpen’s place as a pioneer in Open Access publishing and the home to some of the most relevant scientific research available through Open Access.
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We are so proud to have worked with so many bright minds throughout the years who have helped us spread knowledge through the power of Open Access and we look forward to continuing to support some of the greatest thinkers of our day.
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Thank you for making IntechOpen your place of learning, sharing, and discovery, and here’s to 150 million more!
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Several factors such as natural selection, mutations in genes, the presence of efflux pumps, impermeability of the cell wall, structural changes in enzymes and receptors, biofilm formation, and quorum sensing cause microorganisms to develop resistance against antimicrobials. Isolates that synthesize extended spectrum-β-lactamases (ESBL), induced β-lactamases (IBL), carbapenamases, metallo-β-lactamases (MBLs), and New Delhi metallo-β-lactamases (NDM) have emerged. Determining virulence factors such as biofilms and the level of antimicrobial activities of antimicrobial agents alone and in combination with appropriate doses against microorganisms is very important for the diagnosis, inhibition, and prevention of microbial infection. 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He is an expert in genetic toxicology and is, or has been, a referee for more than 20 international scientific journals. He was a member of the International Panel of Experts at the International Agency for Research on Cancer (IARC, WHO, Lyon, France) in 2015 for the evaluation of DDT, 2,4-D and Lindane. 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She is a member of the National Scientific and Technological Research Council (CONICET) of Argentina in the genetic toxicology field, the Latin American Association of Environmental Mutagenesis, Teratogenesis and Carcinogenesis (ALAMCTA), the Argentinean Society of Toxicology (ATA), the Argentinean Society of Genetics (SAG), the Argentinean Society of Biology (SAB), and the Society of Environmental Toxicology and Chemistry (SETAC). She has authored more than 380 contributions in the field, including scientific publications in peer-reviewed journals and research communications. She has served as a review member for more than 30 scientific international journals. She has been a plenary speaker in scientific conferences and a member of scientific committees. 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1. Introduction
Bats are unique among mammals for their ability to fly. The acquisition of powered flight required a series of morphological and physiological changes in the basic mammal body plan. The structure of the limbs is the most obvious specialization, however, adaptations for powered flight encompass most organ systems, in particular the cardiovascular and respiratory apparatus. Flight performance is strongly determined by wing morphology, which in turn is associated with the biomechanics and energetics of flight, as well as ecological aspects such as foraging behavior and habitat selection.
In this chapter we focus on respiratory, cardiac and wing morphology characteristics of some bat species present in Chile, correlating the results with ecological and behavioral information. The small community of Chilean bat species shows a pattern similar to that found in other bat communities. With respect to wing morphology we found that Tadarida brasiliensis, Desmodus rotundus and Mormopterus kalinowskii have small wing areas, while molossids have high aspect ratios and that of D. rotundus is only moderate. D. rotundus has a smaller mass specific wing span, and the highest wing loading. Myotis chiloensis has a second moment of area of humerus (Ih), lower than expected from allometric predictions, suggesting poorer resistance. Based on these results four functional groups may be recognized: i) species with high wing loading and low wing span such as D. rotundus, capable of rapid flight with moderate power consumption, ii) species with high wing loading and high aspect ratio, such as the molossids T. brasiliensis and M. kalinowski, which are capable of fast flight and low power consumption, characteristic of foragers in open areas; iii) species with low wing loading and low wing span such as most vespertilionids, capable of slow and maneuverable flights in a bat that inhabits wooded areas; and iv) L. cinereus, forming an isolated group characterized by high speed and agility.
Also the respiratory and cardiovascular systems of bats are modifications or refinements that allow them to survive this extreme way of life. Bats have lung volumes about 72% greater than non-flying mammals of the same size. Pulmonary ventilation can rapidly increase 10 to 17 times as flight begins. These respiratory adaptations, along with structural changes of lungs, lead to higher oxygen consumption than other mammals of similar size, reaching up to 22 mlO2/gh at low temperatures and during hovering. We found that the bronchial morphology of T. brasiliensis shows an optimization of the proximal airway with minimum entropy production during mechanical ventilation. In addition, bats have a very thin alveolar-capillary barrier, yielding an oxygen diffusion capacity similar to birds. Also, the heart of bats is larger than in all other mammals, representing about 1% of body weight, reaching in some cases 2%. Birds and bats reach very similar aerobic capacities. However, while birds have a large set of structural changes in their respiratory system, bats have a cardiorespiratory system optimized to their extreme life style.
The order Chiroptera (“winged hands”) is practically defined by saying that it is constituted by flying mammals. These animals require deep structural changes associated with their lifestyle, but based on a mammalian model. Flight influences its main characteristic: wings formed by a membrane called a patagyum. The arms are the dominant limbs while legs are reduced, contributing to the reduction in body mass which is necessary for flight. These structural changes are also associated with the colonization of the crepuscular and nocturnal air space which required the specialization of the visual system in megachiropterans and the development of echolocation in michrochiropterans, where excepting macro chiropterans the vision contributes little, but where the emission and reception of ultrasound, or echolocation, allows the recognition of the surrounding environment; the ear is the main organ sense of the group.
2. Body size
Body size is associated with flight behavior, diet selection, reproductive behavior, physiology and practically all aspects of the biology of bats. (Swartz et al., 2003). Bat body sizes vary from 2 g and 16 cm wingspan in the mammal with the lowest body mass known Craseonycteris thonglongyai, to 1.5 Kg and 2 m wingspan in the Asian flying foxes (Megachiroptera; Pteropodidae) (Fenton, 1992).
The superior limit of body size is not imposed by flight, because among birds there are species which weigh up to 14 Kg, such as Koris`s bustard, and the extinct pterosaurs reached giant sizes. It is possible that in bats the superior limit to body mass is imposed by a combination of behavioral, ecological and physiological factors. Insectivorous bats would have aerodynamic and sensorial restrictions. Barclay & Brigham (1991) proposed that associated with an increase in the body mass there is a decrease in the maneuverability that prey detection at long distances requires. However, this would condition the use of low frequencies during echolocation, with a decrease in spatial resolution. Thus, the abundance of large prey could be a limiting factor of body size in these bats, which is corroborated in part by the positive correlation between prey size and body size of bats (Aldridge & Rautenbach 1987; O\'Neil & Taylor, 1989). However, this does not apply to large fruit bats that do not use echolocation. In the latter restrictions derived from muscle physiology may operate; kinematics of flight or wing loading and mechanical stress imposed on the bones by flight (Marden, 1994). While the force per unit mass generated by a muscle is approximately constant, the mass-specific power to fly scales positively with mass, resulting in less lift generation per unit of muscle power (Marden, 1994). Similarly, the mechanical power required for flight grows faster (α Mb 1.185) than the oxygen consumption of bats (Maina et al., 1991; Maina 2000) helping to establish an upper limit of about 1.5 Kg for bats (Carpenter 1986, Maina 2000).
3. Limbs
Limbs of bats are completely conditioned by flight. While the forelimbs are large and strong, the legs are small, contributing to a reduced mass allowing flying. However, these latter have adaptations such as the joint mechanism of the claws, which pivot on the distal phalanges. While an elastic ligament extends the dorsal claw, the long plantar tendon inserts on the ventral side of the base of the claw, flexing it. Thus, when bats hang inverted during rest, the body weight flexes the claw and allows it to catch on a branch or a cliff (Neuweiler, 2000). In most mammals the diameter of the femur scales with body mass raised to the 1/3 power (geometric similarity), but in bats femur diameter is smaller than that of other mammals of similar size. An exception to this generalization is the vampire bat Desmodus rotundus in which the diameter of the femur follows the curve of non-flying mammals (Swartz, 1997). This species has a semi-quadrupedal locomotion while feeding, being able to travel on all four limbs and even start flight with a jump (Schutt et al., 1997).
The body and forelimbs are significantly modified for flight. The thin patagium is richly vascularized with muscles that allow tension and bending, thus contributing dynamically to flight. Occipitopollicalis muscles, Dorsoplagiopatagialis, Humeropatagialis, Coracocutaneus, Uropatagialis and Plagiopatagial Tensor contribute to this dynamic tension, while the adductor of the fifth digit causes the arched profile necessary for flight. While bird wing movement is controlled mainly by two muscles and the point of rotation of the wing is slightly medial or dorsal to the level of shoulder joint, in the bats this point is shifted ventrally to the sterno –clavicular articulation, allowing the scapula to participate in wing movements. In the movement of bat wings at least 17 muscles are involved (Neuweiler, 2000). The main lift muscles are the Trapezium, rhomboids, Acromiodeltoideus and Spinodeltoideus, while the lowering of the wings is controlled mainly by Pectoralis, Serratus, Clavodeltoid and Subscapularis. Extension and flexion of the wing are governed by a special muscle arrangement that automates these movements. Both the triceps (extensor, dorsal) and the biceps (flexor, ventral) are inserted from the scapula to the forearm, bypassing the humeral insertion. Also the extensor carpi radialis and flexor carpi ulnaris bypass the radius. Thus the contraction of the triceps causes the extension of the radio-carpal extensor and the whole wing in an almost automatic form (Neuweiler, 2000).
There are four important parameters related to the aerodynamics of flight: 1) wing loading:
WL=mg/SE1
which represents the weight per unit area (N/m2) to be supported by the wings; 2) wingspan (B), corresponding to the length of the wings from tip to tip, 3) the aspect ratio:
AR=B2/SE2
,
which is a dimensionless measure of the relative length to width of the wings, so high AR values correspond to long, thin wings and vice versa, and finally 4) wing acuity ratio (i.e., tip length ratio: TL = length of third finger / arm length) (Neuweiler, 2000).
4. Flight
In its most simple terms, a bat must move the air with its wings in such a way as to produce aerodynamic force. The component of the aerodynamic force that propels the bat forward is thrust and the component that keeps the bat from falling is lift. These forces are opposed by drag (an aerodynamic force) and gravity, respectively. In contrast to planes that continuously produce thrust and lift (by their engines and the constant flow over the wings), bats generate aerodynamic force in a cyclic manner due to the flapping of the wings. Thus, flight in bats is dependent of an appropriate modulation of wing kinematics in order to generate enough aerodynamic force.
Unlike terrestrial locomotion, where limbs push against a solid substrate, aerial fliers use their wings to push against fluids, which distort and swirl to form a complex wake (Dickinson et al., 2000). Although it is the wing motion that is directly responsible for the generation of lift and thrust, we can estimate the aerodynamic forces by looking at the fluid motion left behind a flying animal. Newton’s third law requires that the forces exerted by the air upon the wings must be equal and opposite to the forces exerted by the wings upon the air. The wake left behind the wing thus contains a complete ‘footprint’ of its force generation. An everyday example of this are the vapor trails left by airplane wings, the tip vortices, that arise directly from the aerodynamic forces produced as the plane moves through the atmosphere. Bats also leave an aerodynamic wake and this wake can be measured by looking at the movement of the air left behind.
An aerodynamic wake can be efficiently analyzed in terms of its vortex structure. Vorticity is the local angular or rotational velocity of the fluid, and a vortex is somewhat subjectively defined as a concentration of vorticity. Tornados and swirling motions of water draining are familiar examples of vortices. Visualization and quantification of these vortices can be used to estimate aerodynamic forces. Early studies of bat’s wake structures, using helium-filled bubbles, suggested that the upstroke function and wingbeat gaits may vary to flight speed, with lift being produced during upstroke at high speeds but not during slow flights (Rayner et al., 1986). These differences in lift generation would be expressed as discrete vortex rings during slow flight in contrast to the ondulating but constant vortex lines observed at fast flight speeds (figure 1). Recent studies using digital particle image velocimetry (DPIV) have allow us to study the wake left behind flying bats with much higher temporal and spatial resolution than those original studies. The emerging picture of the aerodynamic footprint left by bats is that the wake structures are more complex than expected, potentially because of the more complex wing kinematics than that of birds and insects as well as the compliant characteristics of the wing membrane, and currently it is an area of very active research (e.g., Hedenström et al., 2007; Johansson et al., 2008; Muijres et al., 2008; Hedenström et al., 2009; Hubel et al., 2009; Hubel et al., 2010; Wolf et al., 2010).
Figure 1.
Effect of the oscillation of the wings on the position of the center of mass (COM) and accelerations of the body. When external forces, such as aerodynamic and gravitational forces, are absent, the position of the COM will remain constant but the body moves in opposition to the flapping wings to conserve momentum. Closed and open symbols correspond to the pelvis and chest markers, respectively. During upstroke (A), the upward and backward acceleration of the wings will produce an inertial force (black arrow) that will move the body forward and downward with respect to the downstroke. This force will produce a forward-oriented component, or inertial thrust, during upstroke (grey arrow). During downstroke (B), the downward and forward acceleration of the wings will produce an inertial force (black arrow) that will move the body backward and upward while keeping the position of the COM constant. The horizontal component of this inertial force will produce negative inertial thrust during downstroke (grey arrow).
Aerodynamic theory predicts that the wing loading, the wing span and aspect ratio are significant parameters in determining performance in flight. For example, during flight the organism should generate sustained lift (L) to support body weight and thrust (T) to overcome drag (D). Thus, the power required to fly is:
P=D⋅v=T⋅vE3
,
where v is the relative velocity of air over the wings. The cost of transport (C), which corresponds to the work done to move a unit of weight for a unit of distance is inversely proportional to the speed:
C=P/mgv=T/LE4
,
with P = power, m the body mass and g gravity acceleration). Furthermore, the speed is proportional to WL\n\t\t\t\t1 / 2, so that both high wing loading and high flight speeds are associated with low transportation costs (Norberg 1987).
The energy per unit time (power) required to fly can be decomposed into that needed to move the wings (inertial power: Pin) and the power required to produce the aerodynamic force (R). The latter can be decomposed into the power required to overcome the resistance of the body (parasite power: PPAR), the profile of the wings (Power Profile: PPRO) and the power to generate lift and thrust (induced power: PI). Thus the total aerodynamic power is the sum these:
P=(PPAR+PPRO+PI)+PinE5
.
Plotting the power according to flight speed a typical "U" curve is obtained, whose minimum determines the speed at which it produces the minimum energy expenditure (VMP). It is also possible to calculate the speed which determines the minimum cost of transport (VMR), which is determined by the intersection between the curve and the tangent to it passing through the point (v = 0) (Norberg, 1987).
All these components of energy expenditure of flight are correlated with B and WL, for example Pparα v3 α (WL)3/2, Ppro α S(B/τ)3 during hovering, where τ is the wing beat period, PI α (Mg)3/2/B during hovering and PIα (Mg)2/(B2v) during forward flight, and Pin α B2 (Norberg, 1987). In addition, the minimum resistance (Dmin) and the minimum power required to fly (Pmin) are inversely correlated with the aspect ratio:
Dmin=2mg(Cr/πAR)1/2E6
,
and
Pmin=[0.95(mg)3/2Cr1/4]/B(AR)1/4E7
,
where Cr is the combined parasite and profile friction coefficient. Thus high values of AR are critical in reducing both parameters; AR is considered to be a measure of aerodynamic efficiency (Norberg, 1994). Another important aspect is the high wing acuity (i.e., TL) that allows adequate air movement dynamics around the wings without turbulence. By contrast, rounded wings can generate turbulent flow by increasing the resistance to movement and therefore PPRO.
Norberg and Rayner (1987) attempted to establish a relationship between lifestyle and major aerodynamic parameters of wing morphology, being able to classify four groups of bats: i) bats of open space and faster flight have long and narrow wings, with high wing loadings up 20N/m2 and aspect ratios as high as 14.3 in some Molossidae (Fenton 1992, Norberg & Rayner 1987), ii) slow-flying bats of forested areas with short and broad wings, with low wing loading, about 5 to 6 N/m2, and low aspect ratios, about 5 (Canals et al. 2001, Iriarte-Díaz et al. 2002), iii) fast flying bats with stationary or short flights, which have high wing loading but low aspect ratios, and finally iv) slow-flying bats in open spaces, which have high aspect ratios but low wing loading (Figure 2).
Species that forage in and around foliage tend to have short, rounded wings with low values of AR and TL, which produces low wing loading. They have a relatively slow flight, between 2.5 and 6 m / s, and are very maneuverable (Neuweiler, 2000). Many of them can use hovering to locate and capture prey over the foliage or to feed on pollen or nectar. Species that forage on leaves are slender, with long, thin wings (high AR) and high wing loading. Their flight speed is high, between 9 and 15 m / s. These bats have less maneuverability. However, their agility, defined as the ability to accelerate and stop quickly, is increased, as is an ability related to wing loading (Norberg & Rayner, 1987). An example of such bats is the Molossidae, for example Tadarida brasiliensis in which the highest flight speed has been registered: 27 m / s (Neuweiler, 2000). Some species of bats feed by fishing or hunting on bodies of water, such as Noctilio leporinus, requiring a great generation of thrust (T) in flight, plus an important handling. Thus they have higher TL, AR moderate and relatively low wing loading.
Figure 2.
Principal components for morphological characteristics in several bat species. The first component was explained for body mass, but second and third components are related with wing loading (WL) and the aspect ratio (AR) respectively. This analysis allow recognize different eco-morphological groups of bats. Modified from Wainwright & Reilly 1994.
Frugivorous bats usually fly long distances for foraging, occasionally flying over 27 km. This requires sustained flight and highly developed flight muscles that result in high wing loading, however, their wings are broad and rounded (Neuweiler 2000). The same is true in the vampire bat Desmodus rotundus (Canals et al., 2005). Bats which plane using convection currents such as some Pteropodidae usually have larger wingspan and wing loadings lower than those using flapping flight (Norberg et al., 2000).
4.1. Studies in Chile
In a series of studies, Canals et al (2001), Iriarte et al (2002) and Canals et al (2005) examined some aspects of the wing morphology of 8 species of bats present in Chile, correlating the results with available ecological information. They estimated aspect ratio, wingspan, wing surface, and wing loading of the molossids Mormopterus kalinowskii and Tadarida brasiliensis, the Phyllostomidae Desmodus rotundus and the vespertilionids Myotis chiloensis, Histiotus montanus, Histiotus macrotus, Lasiurus borealis and Lasiurus cinereus (Table 1).
Species
Mb (g)
B (cm)
S (cm2)
WL (N/m2)
AR
Ih (cm4x10-6)
Myotis chiloensis (49)
6.76 ± 0.18
23.69 ± 0.39
98.29 ± 3.47
6.8 ± 0.23
5.76 ± 0.16
3.89 ± 0.49
Histiotus montanus (1)
12.5
29.2
-
-
-
23.1
Histiotus macrotus (3)
9.37 ± 0.29
29.67 ± 0.58
129.67 ± 4.20
7.08 ± 0.19
6.78 ± 0.06
21.17 ± 3.52
Lasiurus borealis (3)
7.87 ± 1.12
25.37 ± 2.49
93.73 ± 8.87
8.20 ± 0.46
6.87 ± 0.70
12.68 ± 9.30
Lasiurus cinereus (2)
19.55 ± 6.58
30.20 ± 1.41
165.45 ± 52.07
15.42 ± 5.75
5.72 ± 1.29
26.78 ± 5.06
Tadarida brasiliensis (27)
11.95 ± 0.62
28.65 ± 0.63
100.14 ± 4.61
11.56 ± 0.66
8.12 ± 0.16
11.15 ± 2.61
Mormopterus kalinowskii (2)
3.10 ± 1.13
17.25 ± 0.35
32.4 ± 2.26
9.28 ± 2.77
9.20 ± 0.27
5.85 ± 5.98
Desmodus rotundus (1)
33.48
33.5
167.23
19.61
6.71
68.3
Histiotus montanus (1)
4.3
19.7
58.8
7.17
6.6
9.3
Lasiurus cinereus (4)
12.23 ± 2.71
27.33 ± 0.68
93.50 ± 8.18
12.74 ± 2.07
8.01 ± 0.44
21.24 ± 14.01
Table 1.
Summary of the aerodynamic characteristics of the wings of eight bats.
* From Iriarte-Díaz & Canals 2002 and Canals et al., 2005.Mb = body mass, B = wing span, S = wing surface, WL = wing loading, AR = aspect ratio and Ih = second moment of humeral area in median section. Asterisks indicate juvenile individuals. In one adult H. montanus it was not possible to estimate the wing surface. Numbers in parentheses indicate sample sizes.
The free-tailed bat T. brasiliensis and D. rotundus have no tail membrane and a low wing area, but while the molossids have high aspect ratios, that of D. rotundus is only moderate. D. rotundus has a smaller wingspan for its body mass, and the highest wing loading. Furthermore, these authors estimated radiographically the second moment of area of humerus (Ih), which corresponds to a measure of bone strength to bending. Myotis chiloensis had a value lower than expected from allometric predictions, suggesting poor resistance. All other vespertilionids showed a high second moment of area, which may be explained by their costly form of locomotion, especially in species with high parasite load as a result of their long ears. The high Ih shown by D. rotundus may be explained by the low aspect ratio and high body mass, which increase the torque produced by the weight during quadrupedal locomotion.
The small community of Chilean bats showed a similar pattern to that found by Norberg and Rayner for many species, but at a small scale. Principal components analysis showed two axes, the first correlated positively with wing loading and negatively with wingspan and the second positively correlated with the aspect ratio. In these species 4 functional groups can be recognized, one for each quadrant in the graph:
D. rotundus, with high wing loading but low wingspan (relative to its body size), located in the high agility and rapid flight zone with moderate power consumption, which is likely related to long flights to their resting places and their particular form of locomotion;
The molossids T. brasiliensis and M. Kalinowski in the area of high flight speed and low power consumption as is characteristic of high speed open area foragers;
Most of vespertilionids in the zone of high maneuverability and low speed which correspond to bats which inhabit wooded areas;
L. cinereus forming an isolated group in a zone of high speed and agility.
4.2. The mechanics of flight: Kinematics
The differences in flight performance observed in bats can be associated with higher energy expenditure efficiency as well as very high levels of maneuverability. For example, among animals of comparable body size, hovering flight of nectar-feeding bats is 40 and 60% less costly metabolically that that of hawkmoths and hummingbirds, respectively (Winter, 1998; Winter and von Helversen, 1998; Voigt and Winter, 1999), suggesting that bats have more efficient mechanisms of lift generation than member of other groups. Although the kinematics of hovering of bats differ from those of insects and hummingbirds, we lack experimental measurements that can explain such differences in efficiency. In a recent study using PIV methods, it was shown that bats can increase lift generation during slow flights by 40% by using attached leading-edge vortices around the wings (Muijres et al., 2008), similar to those used by insects (Fry et al., 2005) and hummingbirds (Warrick et al., 2005) during hovering flight. Why hovering flight in bats is energetically cheaper than that of insects and hummingbirds of similar size is still unclear.
4.3. Maneuvering during flight
The ability to quickly alter flight direction and speed is essential for bats to successfully navigate complex three-dimensional environments, to capture prey, and to avoid predators. Despite the importance of this task, maneuvering abilities and its mechanisms have been barely investigated. A flying organism has six degrees-of-freedom of movement: translation in three dimensions in space and rotation around three orthogonal axes centered on the center of mass, termed yaw, pitch, and roll.
In its most basic form, a turning maneuver requires the reorientation of the body in such a way that the net aerodynamic force is tilted laterally effectively producing a centripetal force that will drive the bat through the turn. The most common method in the literature is the bank turn. In this kind of turn, the body rolls into a bank, which orients the lift vector towards the direction of the turn, producing a centripetal force. When the turn is complete, the body rolls back into the unbanked position such that centripetal force is no longer produced. Airplanes use this mechanism, it has been observed in insects and birds, and it has been assumed that bats use it as well. If a flying organism performs a banked turn, then for any given lift coefficient and bank angle, the turning radius depends directly of the wing loading or body weight per unit wing area; there is some evidence consistent with this relationship from bats in both field and obstacle course settings (Aldridge, 1986; Aldridge and Rautenbach, 1987; Stockwell, 2001).
However, growing evidence suggest that differences in turning techniques (e.g., gliding versus flapping turns, Aldridge, 1987b) and changes in wing posture throughout the turn (Lentink et al., 2007) can substantially alter the turning performance. The only study to investigate the mechanisms of turning in bats suggest a more complex mechanism. Detailed analysis of the wing motion and body orientation during 90-degree turns in the pteropodid Cynopterus brachyotis showed that during the upstroke the body rotates into the direction of the turn, a mix of roll and yaw rotations, without changes in flight direction. This body rotation allows the bat to use part of the thrust generated during the downstroke to enhance the centripetal force from the bank turn, thus allowing the bat to perform tighter turns than predicted by wing morphology alone (Iriarte-Díaz and Swartz, 2008). These results highlights the importance of studying the mechanics flight performance and that using morphological proxies to estimate performance (e.g., wing loading and aspect ratio) might severely underestimate flight abilities of bats.
4.4. The effect of wing inertia during flight
One aspect of flight performance that remains virtually unstudied is the importance of inertial forces generated by the flapping motion of relatively massive wings. The wings of bats comprise a significant portion of total body mass, ranging from 11 to 20% in a few measured species (Thollesson and Norberg, 1991; Watts et al., 2001) and consequently, inertial forces produced by accelerating these masses are expected to be high and the potential effect of these forces on flight performance is still not well understood. In a recent study, the effect of wing’s inertial forces was studied on C. brachyotis during forward, steady flight (Iriarte-Díaz et al., 2011 in press). At any speed, the tip of the wings move upwards and backwards relatively to the body, but if the speed of the body is low enough, the tip can sometimes move backwards relatively to the still air during the upstroke. This backward movement of the wingtip has been called “tip-reversal upstroke” (Aldridge, 1987a) and for decades has been thought that it provides additional thrust to slow-flying bats, partially because the observation that the bat’s trunk accelerate forward during upstroke. However, for C. brachyotis, the forward acceleration of the body is the result of forwardly directed inertial forces produced by the motion of the wings. When the wings swing backwards, approximately 20% of the bat’s mass moves backward relative to the center of mass. In order to maintain the momentum, other portions of the body must move forward relative to the center of mass, which is reflected in the forward acceleration of the trunk. Using a model of the mass distribution of the trunk and wings, inertial accelerations were estimated and removed in order to estimate the acceleration of the center of mass. When inertial forces were removed, forward acceleration of the center of mass only occurred during the downstroke (Iriarte-Díaz et al., 2011). Thus, inertial forces may be potentially important during flight, although when and how they can be used is not known. Preliminary evidence suggest that inertial forces might be important during turning (Iriarte-Díaz and Swartz, 2008) and when performing landing maneuvers (Riskin et al., 2009).
5. Physiology and energy of bats
The first law of thermodynamics states that energy is neither created nor destroyed, only transformed. Living organisms as physical systems obey this principle, acquiring, converting, assigning, storing and dissipating energy. The transformation of energy plays a crucial role in the evolution, ecology and physiology of organisms. Thus the internal and external boundaries of the use and transformation of energy affect their fitness and may affect species richness, reproductive effort, activity patterns, habitat use and life history (McNab 2002, Cruz-Neto et al 2003; 2006).
Field metabolic rate (FMR) integrates all the energy costs of free-living organisms, including the costs of thermoregulation, locomotion etc. This has been quantified in the Australian bat Syconycteris australis with doubly labeled water, reaching 7 times the basal metabolic rate (Geiser & Coburn 1999), one of the highest values described in endotherms. This is attributable to the prolonged nocturnal flight (Geiser 2006). The mass specific metabolism of bats is 1.6 times that of non-flying mammals (Thomas, 1987). Also, during flight metabolism increases to 20 or 30 times the standard rate (Thomas, 1975). During flight, about 25% of metabolism is converted into work, so that 75% is dissipated as heat. This is done primarily in two ways: via airway and skin. The airway dissipates only about 15% because little or no frequency change is possible during flight, since there is a synchrony between wing beat and respiration. Thus the skin must remove the remaining heat (85%). Its large area and conductance allow this removal of heat by convection and radiation, which is favored by vasodilatation and opening of arteriovenous shunts in the wings and by the greater thermal difference between the body and the environment during nocturnal flying.
The thermal conductance of a microchiropteran of 10 g is about 6 times that of a mega-chiropteran of 500 g (Geiser 2006). Thus the maintenance of homeothermy is especially relevant in small bats with large membranous wings and large lungs. Bats have a respiratory area 6 times greater and a conductance between 1.5 and 4 times greater than non-flying mammals (Neuweiler, 2000), although the minimum conductance at rest appears to be similar (see Speakman & Thomas 2003). Despite this, bats can remain active and euthermic within wide temperature ranges.
To maintain their temperature bats may use different behavioral and physiological strategies. Behaviorally they can avoid overheating by wing movements that favor convection or licking the surface of their skin to increase evaporation, since they do not have sweat glands. Small bats find microenvironments with high thermal stability in caves or shelters, and can travel to other shelters to avoid overheating at times of high temperatures. Thus, the solitary bat Syconycteris australis resting under leaves selected thermal environments in the middle of wooded patches in spring and autumn, protected from the extreme temperatures, while in winter it moved to the extremes (Law 1993). To avoid cold behaviorally, many species have a social grouping behavior (huddling) (Roveroud & Chappel 1991), or nest in caves forming large colonies of hundreds to millions of individuals, raising the temperature up to 8° C above that of the rest of the cave (Dwyer & Harris 1972).
The first physiological response to cold is to increase muscle tone generating heat, followed by shivering, which actually consists of rhythmic but asynchronous fibrillary muscle contractions. However, heat generation consumes so much energy that with limited resources it is not convenient for long periods.
When ambient temperatures fall below the lower limit of thermoneutrality, bats have the "option" to maintain their body temperature at a high energy cost, or to enter into torpor, maintaining a temperature similar to that of the environment with a significant decrease in energy expenditure. Entering into torpor seems to depend upon the interaction among resource availability, reproductive status and body size. McNab (1983) proposed a boundary line of endothermy, allometrically related to body size with an exponent -0.67, which intersects the Kleiber line for mass-specific metabolism at 37 g. Thus, individuals under this line and weighing less than 37 g may use torpor as a physiological response to save energy. During torpor, animals enter into a rhythmic pattern of breathing and apnea; the periods are longer as the temperature drops.. These periods allow the accumulation of CO2 that triggers breathing and prevents evaporation in the lungs. The mechanism that triggers the awakening is still unknown. Many bats have facultative stupor, maintaining significant fluctuations of oxygen consumption and body temperature (heterothermy), saving a great amount of energy. Some examples of this behavior are found in Eptesicus fuscus, Rhinolophus ferrunequinum, bechteinii Myotis, Myotis evotis, Lasiurus cinereus (Willis 2006) and Myotis chiloensis (Bozinovic et al. 1985). These bats have different patterns, such as the presence of lethargy in the daily rhythm with no difference between sexes, preference in pregnant and breast-feeding and preference for males. These depend on the fine balance between the costs of endothermy, reproduction and locomotion costs imposed by the high wing loading in the pregnant females. For example, if the costs are very high, some bats prefer to rest in cold and go into torpor, avoiding the cost of endothermy and negative energy balance (Willis 2006).
Another long-term mechanism for energy saving is hibernation, in which metabolism falls to extremely low levels; heart rate can also decrease from more than 400 beats to a few beats per minute and the peripheral circulation and urine output may fall to almost nil. The respiratory quotient drops to 0.6-0.7, indicating that metabolism of lipids and blood glucose values may reach about 25 mg / dl. In contrast to torpor in which the values of Q10 (ratio of metabolic change with 10° C of temperature change) are around 2, during hibernation they are temperature-dependent, increasing from 2 to 4 with an increase of temperature, which suggests an active metabolic depression. For example at 20° C the metabolism of a bat in torpor is twice that of a hibernating bat.
This metabolic depression could be due to metabolic acidosis, low thyroid hormone or mediated by fatty acids (Neuweiler, 2000). The mechanisms that trigger hibernation have not been established, although it has been postulated that hibernation is regulated autonomously. Temperature, energy depletion and loss of water have been postulated as triggers that regulate arousal.
5.1. Energy balance: Myotis chiloensis, a case study
The perpetuation of animals over time requires an average positive energy balance, which is particularly difficult for small mammals such as the insectivorous bat Myotis chiloensis. Bozinovic et al (1985) studied the oxygen consumption of 25 individuals (5.78 ± 0.9g) at different temperatures. There were two responses: a) euthermic metabolic levels (36.6 ± 2.2° C) with an average oxygen consumption of 1.76 mlO2/gh b) torpor metabolic levels, where the temperature was only 0.5° C above room temperature with metabolic reductions of 81 to 98% (Figure 3). In continuous records M. chiloensis showed a daily rhythm with only three hours in which animals were euthermic.
The torpor in M. chiloensis was as expected for a small mammal, under the endothermy limit proposed by McNab (1983); however it does not explain why this species does not maintain high metabolism all the time.
The answer seems to come from the energy balance
Intake−Egestion=Mactivity+Meuthermia+Mtorpor+EE8
,
where M is metabolism and E the energy balance.
Figure 3.
A) Relation between ambient temperature and metabolism (MR, mlO2/gh) and B) Relation between ambient temperature and body (Tb, °C) in Myotis chiloensis in euthermic state (black circles) and in torpor (white circles) (Modified from Bozinovic et al., 1985).
Data on the chemical composition of various flying insects indicates that their assimilated energy is approximately 5.3 Kcal / g; thus an individual of 5.8 g which ingested 11% of its weight in insects every day assimilates 5.8 x0, 11x5, 32 = 3.39 kcal / day. Therefore this individual may have two situations:
Euthermy: The temperature the shelters of Myotis chiloensis averages 19.5° C and metabolism at this temperature is 36 cal / gh, so that in the 21 hours of resting the minimal euthermic energy expenditure is 5.8 x36x21 = 4.38 kcal. The rest of the time (3 hours) M. chiloensis is flying and feeding. Assuming that the metabolic activity is at least three times BMR, the energetic cost would be 5.8 x36x3x3 = 1.88 kcal. Thus we have the following relationship: 3.39 = 4.38 + 1.88 + E, where the euthermic energy budget would be E = -2.87 kcal / day, i.e., the individual would have an energy imbalance equivalent to a mass loss of 5 to 10% per day.
Torpor: If instead it spends 20 hours in torpor with a metabolism of 1.2 cal / gh, 1 hour of euthermic rest (30 min before and after feeding) and the same three hours of activity, it would expend 5.8 x1, 2x20 = 0.14 Kcal in torpor, the same 1.88 Kcal during activity and 1x36x5, 8 = 0.21 Kcal at euthermic rest, giving a balance 3.39 = 0.21 + 1.88 + 0.14 + E. Now the energy budget is positive: E = +1.16 kcal / day.
6. The respiratory system
Endothermic animals depend on aerobic metabolism for most of their vital functions. The energy from food is allocated to different functions such as maintenance of homeostasis (i.e. temperature), reproduction, exchange mechanisms, maintenance of tone and locomotion. As most bats are small and therefore have a large surface area per unit volume, they have trouble maintaining their body temperature high and constant as consequence of the significant energy loss through the skin. Moreover, flight requires high energy expenditure, especially since many bats are exposed to cold nights and fly at high altitudes with low oxygen partial pressures (Harrison & Roberts 2000). In this sense bats may be considered as mammals adapted to extreme environments where oxygen management is crucial. Both the respiratory and cardiovascular systems undergo changes or refinements that allow them to optimize the acquisition and delivery of oxygen to tissues, and thus survive this extreme way of life.
Breathing in mammals consists basically of two connected events: ventilator convection and alveolar diffusion. The first is the displacement of a volume of air through the airway and the second in the effective exchange of oxygen and CO2 at the alveolar level.
7. Ventilator convection
Alveolar ventilation (V˙) may be expressed as the product of effective tidal volume (tidal volume (Vc) minus dead space (E)) and the respiratory rate (fr):
V˙=(Vc−E)⋅frE9
.
Thus increments in ventilation are possible only through effective tidal volume or respiratory rate increments. However, increasing alveolar ventilation may be costly in energetic terms as the movement of larger volumes of air results in greater breathing work. Moreover, the work of breathing (Tr) not only depends on the volume but also on the pressure necessary to mobilize this volume:
Tr=∫PdVE10
.
This in turn is a direct function of the resistance to air movement which is influenced by a) a geometric factor:
Rg=8ηlπr4E11
,
(Poiseuille Law), where l is the length of the airway, η the air viscosity and r the radius of the bronchi, which basically indicates that the resistance to flow is inversely proportional to the fourth power of the radius, and b) a dynamic factor:
Rv=k1⋅v+k2⋅v2E12
,
where v is the velocity of air flow, which indicates higher resistance at higher flow rates (or breathing rates). Thus the total resistance to airflow through the airway as a function of breath rate follows a U-shaped curve, determining for each species, according to the geometric characteristics of the airway, an optimal respiratory rate with minimal resistance. Murray (1926) and later Weibel and Gomez (1962) and Wilson (1967) established that respiratory geometry could be adapted to a minimum overall work of breathing and minimum entropy dissipation during mechanical ventilation, following approximately the Murray law "For minimum breathing work, ventilation (Q: minute volume) should be proportional to the third power of the radius (r):
Q=k⋅r3E13
.
However, mammals have considerable deviations from this pattern, especially due to the presence of asymmetries in diameter in the bronchial bifurcations and non-uniform length of segmental and subsegmental bronchi (Horstfield, 1990, Canals et al., 2002).
Bats have a much greater lung volume than non-flying mammals and they remove about 60% of the total lung capacity with each breath during flight (Neuweiler, 2000). Lung volume is about 72% greater than in non-flying mammals of similar weight (Canals et al, 2005a) (Table 2). At rest, pulmonary ventilation is similar to that of non-flying mammals. However, this can rapidly increase 10 to 17 times when flight begins (Thomas, 1987). This is due to increases of 3 to 5 times in breath rates and 2 to 4 times in tidal volume. The respiratory rate is synchronized to the wing beat frequency, reaching a value of 400 min-1. These respiratory adaptations function together with structural changes of lung yield in oxygen consumption reaching to 2.5 to 3 times higher than mammals of equal size (Thomas, 1987) and high maximum oxygen consumption, which can reach 22 to mlO2/gh at low temperatures (Canals et al., 2005b) and during hovering (Winter et al., 1998, Voigt & Winter, 1999; Voigt, 2004).
The morphology of the airways also appears to play a role in saving energy during flight. Canals et al. (2005) studied the airway of Tadarida brasiliensis, finding that this species showed fine adjustments in the geometry of the bronchial bifurcations leading to a better optimization of the proximal airway. As the airway is responsible for 80% of lung resistance and 80% of this is generated in the proximal airway, the optimization of the proximal airway can mean less energy loss during flight (Figure 4).
Optimization of the proximal airway of Tadarida brasiliensis. T. brasiliensis is compared with the rodents Abrothrix olivaceus and A. andinus. The ordinate is the distance from the optimum value determined by the geometry of the bronchial bifurcations. While rodents have values farther from the optimum, T. brasiliensis shows a better optimization in the proximal zone, which is the key to a reduction in respiratory work, dissipating less energy. Different letters represent statistically significant differences
8. Alveolar diffusion
The diffusion of oxygen through the alveolar-capillary barrier depends directly on the gradient of partial pressure of oxygen between the alveoli and the capillary (ΔPO2) and the respiratory surface (A), and inversely on the thickness of the alveolar-capillary membrane (τh) This can be expressed as:
V˙O2=κdSa⋅Vpτh⋅ΔPO2E14
,
where the alveolar surface is expressed as the product of lung volume (Vp) and the surface density per unit of lung volume (dSa), κ is Krogh\'s constant and VO2 is the oxygen consumption (Weibel et al., 1981). Thus, high oxygen consumption may be achieved through increases in alveolar surface density or lung volume, or by reducing the thickness of the alveolar-capillary barrier. The factor:
DO2=κdSa⋅VpτhE15
,
is known as conductance or oxygen diffusing capacity (DO2). As mentioned above, bats have a lung volume 1.72 to 1.75 times that of non-flying mammals, however, alveolar surface density is similar to that of non-flying mammals (Maina, 2000). As a result, the total respiratory area of bats is larger than in non-flying mammals. In addition, these animals have a very thin alveolar-capillary barrier (Maina et al., 1991; Maina, 2000a) that may reach a value of 0.1204 microns in Phyllostomus hastatus, the lowest measured in mammals. So bats have very high oxygen diffusion capacity, similar to those of birds (Table 3).
9. The cardiovascular system
Respiratory adaptations are insufficient to ensure adequate oxygen delivery to tissues, so these must be accompanied by changes in the cardiovascular system. Here the blood flow generated by the heartbeat, the resistance to flow, and transport of oxygen in the blood are all relevant.
9.1. The heart
Blood flow (Q) can be expressed as the product of volume ejected in each beat (VE) and heart rate (fc) or as the ratio between the gradient of pressure to generate the flow (ΔP) and peripheral resistance (R):
Q=VE⋅fc=ΔP/RE16
.
Peripheral resistance follows a Poiseuille relationship and cardiac work, similar to respiratory work, depends on expulsive volume and pressure:
Tc=∫PdVE17
.
Thus high flow is obtained by increasing the expulsive volume or heart rate and by decreasing peripheral resistance.
Rodents
Mb (g)
τh (μm)
Dsa (cm-1)
VLp (cm3)
DtO2/Mb (mlO2 s-1 Pa-1 g-1 )
Abrothrix olivaceus
26.3 ± 2.00
0.303 ± 0.037
589.37 ± 45.26
0.846 ± 0.28
2.565x10-6
Abrothrix andinus
25.48 ± 1.93
0.345 ± 0.057
791.8 ± 229.68
0.972 ± 0.12
3.589x10-6
Phyllotis darwini
75.0 ± 4.96
0.223 ± 0.033
1140.5 ± 92.0
1.91 ± 0.17
4.92 x10-6
Bats
Tadarida brasiliensis
11.25 ± 0.50
0.230 ± 0.086
690.28 ± 156.96
0.585 ± 0.09
6.398x10-6
Myotis chiloensis
6 ± 0.10
0.219 ± 0.015
2020.3 ± 71.0
0.360 ± 0.01
20.4x10-6
Birds
Zenaida auriculata
142 ± 1.55
0.171 ± 0.026
3102.9 ± 175
3.77 ± 0.06
9.28x10-6
Columbina picui
39.9± 1.4
0.302 ±0.118
2328.9 ±426.4
1.04± 0.04
4.19x10-6
Metropelia melanoptera
78.4± 2.4
0.186± 0.008
2580.4± 190.3
2.11 ±0.07
7.04 x10-6
Notoprocta predicaría
398 ± 11.7
0.469 ± 0.019
1811.3 ± 27
11.32 ± 0.43
2.07x10-6
Table 3.
Pulmonary parameters of some Chilean species rodents, bats and birds. Mb = body mass; τh6 = harmonic mean of alveolo-capillary barrier thickness; Dsa = density of respiratory surface; VLp = volume of lung parenchyma and DtO2/Mb = mass-specific oxygen diffusion capacity of the alveolo-capillary barrier (data from Canals et al., 2005b; Figueroa et al., 2006; Alfaro et al., 2010).
Bats have the largest hearts of mammals relative to body mass, usually representing about 1% of body weight (Neuweiler, 2000), but reaching 2% (Jurgens et al., 1981, Canals et al., 2005a) (Table 4). They have great development of the right ventricle associated with better lung perfusion and high density of capillaries per unit volume. They also have the highest level of energy reserves in the form of ATP that has been measured in the heart of any animal (Neuweiler, 2000). Despite increased cardiac output, the volume expelled is similar to other mammals, somewhat greater than 1.5 ml / kg, indicating that the increase in heart size is mainly at the expense of muscle hypertrophy. The heart rate is extremely variable and may range from a few beats per minute during hibernation to over 1000 beats per minute during flight (Wolf & Bogdanowics, 1987, Neuweiler, 2000).
Species
Mb (g)
Mh (g)
RHM=Mh/Mb (%)
Mh obs Mh exp
Tadarida brasiliensis
11.25 ± 1.13
0.145 ±0.033
1.29 ± 0.23
0.943 ± 0.176
Mormopterus kalinowski
3.1 ± 1.13
0.057 ± 0.018
1.88 ± 0.10
1.041 ± 0.022
Myotis chiloensis
6.88 ± 0.47
0.096 ± 0.0145
1.40 ± 0.20
0.921 ± 0.137
Histiotus macrotus
9.65 ± 0.61
0.166 ± 0.0350
1.71 ± 0.03
1.213 ± 0.237
Histiotus montanus
12.5
0.272
2.18
1.627 ± 0
Lasiurus borealis
7.87 ± 1.10
0.120 ± 0.02
1.55 ± 0.27
1.046 ± 0.169
Lasiurus cinereus
12.76 ± 2.74
0.173 ± 0.042
1.40 ± 0.04
1.042 ± 0.279
Pipistrellus pipistrellus *
4.85 ± 0.18
–
1.26 ± 0.24
–
Myotis myotis *
20.6 ± 0.9
–
0.98 ± 0.08
–
Molossus ater *
38.2 ± 1.4
–
0.97 ± 0.01
–
Phyllostomus discolor *
45.2 ± 1.34
–
0.94 ± 0.09
–
Rousettus aegyptiacus *
146.0 ± 7.5
–
0.84 ±0.08
–
Table 4.
Heart size of several bat species. Mb = body mass; Mh = heart mass; RHM = relative heart mass; Mhobs/Mhexp = ratio of observed to that expected by allometry. (Data from Canals et al., 2005a; Jurgens et al., 1981*)
9.2. Vessels, the resistance to flow and oxygen transport in blood
The vessels of bats follow a mammalian pattern, with some arterial and venous modifications. Unlike other mammals, the venous return of the forelimbs occurs through two vena cava; inferior vena cava have a muscular zone that allow regulation of venous return, lower during rest and high during flight. The arteries of the wing branch into arterioles with a muscle base which can regulate the flow to the wings and maintain the arteriovenous differential pressure. There are also arteriovenous shunts and venous vessels with pulsating zones (venous hearts) that can regulate the return of blood from the wings. The volume of blood is similar to other mammals as well as the affinity of hemoglobin. However, bats have the highest levels of hematocrit measured in mammals and may reach values above 70% in Tadarida brasiliensis and Miniopterus minor. Red blood cells are smaller (Figueroa et al., 2007) and hemoglobin has been found in higher concentrations (18-24 g/100ml blood), similar to that found in hummingbirds (Johansen et al., 1987). Consequently, bats have a transport capacity of oxygen in the blood of 25 to 30%. In comparison, non-flying mammals have an oxygen-carrying capacity of about 18% (Thomas, 1987; Neuweiler, 2000).
10. The narrow-based, high keyed strategy
By comparing the structural and functional adaptations in birds and bats it can be established that they reach very similar aerobic capacities. However, strategies to achieve these high performances during flight are different. Birds have a large set of structural changes in their respiratory system, such as air bags, parabronchi systems, respiratory capillaries, cross-current flows, etc. In contrast, bats have a cardio-respiratory system fully modified to accomplish an extreme way of life. This mammalian structural base is highly refined, operating near maximum values (Maina, 1998) (Table 5). Thus, Maina (1998) comparing a set of 7 parameters including birds, bats and non-flying mammals, found that bats have higher "degrees" of optimization in 5 of them: resting respiratory rate, hematocrit, hemoglobin concentration, resting heart rate and blood count.
Optimization Strategy
Respiratory Adaptations
Cardiovascular Adaptations
Increase of lung volumen
Increase of heart size
Thin alveolo-capillary membrane
Development of right half of the heart
Small alveoles
Regulation of venous return
High oxygen diffusing capacity
High hematocrit
High respiratory frequency
Small GR
Proximal airway adjusted to lower energy loss
Greater concentration of hemoglobin
Greater oxygen transport capacity
Table 5.
Strategy of respiratory and cardiovascular optimization in bats.
Acknowledgments
We thanks FONDECYT grants 100673, 1040649, 1080038 and 1110058.
\n',keywords:null,chapterPDFUrl:"https://cdn.intechopen.com/pdfs/19662.pdf",chapterXML:"https://mts.intechopen.com/source/xml/19662.xml",downloadPdfUrl:"/chapter/pdf-download/19662",previewPdfUrl:"/chapter/pdf-preview/19662",totalDownloads:3691,totalViews:1477,totalCrossrefCites:2,totalDimensionsCites:6,hasAltmetrics:1,dateSubmitted:"November 28th 2010",dateReviewed:"April 19th 2011",datePrePublished:null,datePublished:"September 9th 2011",dateFinished:null,readingETA:"0",abstract:null,reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/19662",risUrl:"/chapter/ris/19662",book:{slug:"biomechanics-in-applications"},signatures:"Mauricio Canals L, Jose Iriarte-Diaz and Bruno Grossi",authors:[{id:"52742",title:"Dr.",name:"Mauricio",middleName:null,surname:"Canals",fullName:"Mauricio Canals",slug:"mauricio-canals",email:"mcanals@uchile.cl",position:null,institution:null},{id:"54276",title:"Dr.",name:"Jose",middleName:null,surname:"Iriarte-Diaz",fullName:"Jose Iriarte-Diaz",slug:"jose-iriarte-diaz",email:"josdiiri@gmail.com",position:null,institution:{name:"University of Chicago",institutionURL:null,country:{name:"United States of America"}}},{id:"54277",title:"Dr.",name:"Bruno",middleName:null,surname:"Grossi",fullName:"Bruno Grossi",slug:"bruno-grossi",email:"bgrossi@uab.cl",position:null,institution:null}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Body size",level:"1"},{id:"sec_3",title:"3. Limbs",level:"1"},{id:"sec_4",title:"4. Flight",level:"1"},{id:"sec_4_2",title:"4.1. Studies in Chile ",level:"2"},{id:"sec_5_2",title:"4.2. The mechanics of flight: Kinematics",level:"2"},{id:"sec_6_2",title:"4.3. Maneuvering during flight",level:"2"},{id:"sec_7_2",title:"4.4. The effect of wing inertia during flight",level:"2"},{id:"sec_9",title:"5. Physiology and energy of bats",level:"1"},{id:"sec_9_2",title:"5.1. Energy balance: Myotis chiloensis, a case study",level:"2"},{id:"sec_11",title:"6. The respiratory system",level:"1"},{id:"sec_12",title:"7. Ventilator convection",level:"1"},{id:"sec_13",title:"8. Alveolar diffusion",level:"1"},{id:"sec_14",title:"9. The cardiovascular system",level:"1"},{id:"sec_14_2",title:"9.1. The heart",level:"2"},{id:"sec_15_2",title:"9.2. Vessels, the resistance to flow and oxygen transport in blood",level:"2"},{id:"sec_17",title:"10. 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O.1998Gas exchange during hovering flight in a nectar-feeding bat Glossophaga soricinaJournal of Experimental Biology201237244'},{id:"B77",body:'WinterY.1998Energetic cost of hovering flight in a nectar-feeding bat measured with fast-response respirometry.Journal of Comparative PhysiologyB 168434444'},{id:"B78",body:'WinterY.VonHelversen. O.1998The energy cost of flight: do small bats fly more cheaply than birds? Journal of Comparative Physiology B\n\t\t\t\t\t168105111\n\t\t\t'},{id:"B79",body:'WolfM.JohanssonL. C.VonBusse. R.WinterY.HedenstromA.2010Kinematics of flight and the relationship to the vortex wake of a Pallas’ long tongued bat (Glossophaga soricina)Journal of Experimental Biology21321422153'},{id:"B80",body:'WolkE.BogdanowicsW.1987Hematology of the hibernating bat, Myotis daubentoni.Comparative Biochemistry and Physiology 88A, 63739'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Mauricio Canals L",address:"",affiliation:'
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1. Introduction
Heavy metals, the name has so many definitions based on various parameters. Based on density the metals which are having a density values greater than 5 g/cm3 are considered as heavy metals [1]. According to this study, the heavy metals which would consider as most threat to human beings are lead, cadmium, mercury, and arsenic. Duffs [2] reviewed the usage of the term heavy metals from the history and finally, he concluded that using the term “heavy metals” is meaningless. He established that there is no relation between the density of the metal and to the usage of the term. In the case of heavy metals, metalloid arsenic also included, from this the term heaviness means may be toxicity.
Some of the heavy metals are having so much of biological importance in trace amounts [3] particularly the elements that are present in the 4th period in the modern periodic table. The biological importance of these metals is enzyme functioning (vanadium and manganese), hormone functioning, production (selenium), cellular growth (nickel), and metabolic growth (arsenic). But these metals are required for the human in trace amounts only if their amount in the body increases they cause adverse effects on human health. Overall the heavy metal should be considered as having high density and also biological importance in trace amounts.
There is a lot of importance for the determination of heavy metals in the various environmental segments, such as air, water, and soil due to their carcinogenic and toxic nature. The IARC (International Agency for Research on Cancer) declared arsenic, hexavalent chromium, cadmium, and nickel and their compounds as group 1 carcinogens (proven carcinogens). Arsenic and their compounds cause urinary bladder, liver, and lung cancers. Hexavalent chromium causes lung cancer and nickel and its compounds cause nasal cavity and lung cancers. All these elements cause cancers to human beings by the route of exposure is through inhalation and ingestion [4]. Regarding the availability of various heavy metals in the earth’s crust is about 5%, among which iron occupies nearly 95% [5].
Due to their toxicity and carcinogenic nature, most of the researchers all over the world are reported about the determination and health implications of heavy metals in the environment. Some of them are discussed hereunder.
Jyothi and Mohamed Farook [6] reported the sources, exposure, and toxicity of mercury. [7] reviewed the heavy metal concentrations in ambient air. This study is limited to the estimation of toxicity of heavy metals in the Indian atmosphere and another study [8] they reviewed the heavy metal determination in ambient air all over the world. Kim et al. [9] reviewed heavy metal toxicity and chelating therapeutic strategies. Giller et al. [10] reviewed the toxicity of heavy metals in microorganisms in agricultural soils. This study found that the microorganisms in soil are much sensitive to heavy metal toxicity than animals and plants. Yabe et al. [11] summarized heavy metal pollution and its impact on the environment and the human population in Africa. Das et al. [12] reviewed the toxicity of cadmium in plants. Proshad et al. [13] reviewed the toxicity of heavy metals in soils of Bangladesh. In this study, they concentrated on the impact of industrialization on the concentration of heavy metals in soil. Su et al. [14] reported the heavy metal contamination in soil worldwide. In this study, they mentioned the current situation of contamination and remediation methods.
The present chapter describes the heavy metal sources, exposure, and the impact of their toxicity on various environmental segments. Based on the toxicity and non- biodegradable nature of lead, cadmium, mercury, and arsenic, the present study mainly focused on these metals.
2. Types of heavy metals
Based on the survey of literature the metals that are considered as heavy metals are chromium, lead, cadmium, iron arsenic, cobalt, mercury, copper and zinc are the Heavy metal. According to Kim et al. [9] studies heavy metals have been classified in to two types as essential and non- essential (Table 1). Essential Heavy metals are less toxic at low concentrations and they act as coenzyme in biological process. For example Hemoglobin and Myoglobin consist of Iron, Vitamin B12 consist of cobalt. Non-essential heavy metals are highly toxic even at very low concentrations, they are non -biodegradable and cause severe toxic effects to living organisms.
2.1 Heavy metal toxicity
Some heavy metals are essential to the human biological process, but depending upon their dosage intake leads some unexpected hazardous effects on health and the physiological system. According to [9] studies shows that despite of its beneficiary health effects, heavy metals are acting as carcinogenic agents. Dissolved forms of these metals through different forms as soil pollutants, water pollutant and air pollutants entering into food chain and finally ending in humans, these are leading to severe damage to the cellular system and leading to expose towards cancer. According to the reports of the International agency for research on cancer non-essential heavy metals (As, Cd, Cr) are major cancer- causing agents [9].
2.2 Sources, exposure, and environmental impacts of lead
The sources of lead varies with different countries based on old and new usage of lead products. It is not limited to the processing of gold ore and recycling of used lead products. It is found that the decrease in blood lead levels in the population of the countries in which unleaded gasoline is in usage [15].
A recent study [16] has reported elevated blood levels in pregnant women in a rural village in Bangladesh. In this study, they found more than 30% of women they sampled were had lead levels in blood in the range of above 5 μg/dL. They found the major source of lead exposure to these women were identified as food storage cans. Nearly 18% of food storage cans (out of the tested) were having lead soldering insides and are responsible for lead contamination in these women. Another study in China [17] determined the blood lead levels (BLL’s) in children who are taking treatment in lead specialty clinics. In this study, they found the BLL’s ranging from 5 to 126 μg/dL. The major reasons they found for the higher lead levels in their blood as industrial sources and folk medicine which is popular in China. Another important thing was determined as it is difficult to find lead poisoning in children due to non-specific symptoms. A very recent study from Australia [18] determined the higher lead levels in children due to the high concentration of lead in soil and pretty dish dust at their premises. This study found that the population who are living in old houses built before 1940 are diagnosed with higher lead levels due to pretty dish dust.
In Nigeria, lead poisoning in the population was observed in the area of Zamfara state which contains gold mining activities. Mahuta [19] reported that in Nigeria, the sources of the lead include mining of gold, lead pipes used for drinking water, and cultural usage of lead.
Based on the studies in all parts of the world it is assumed that the sources of lead are historical usage of lead, industrial activities, and leaded gasoline. Major studies reported that children are the most common victims of lead poisoning. The way of exposure includes the inhalation through the nose and ingestion through drinking water and soil.
There are several ways to minimize the lead levels in the environment such as remediation techniques (in soil), using adsorbents (in water), and using unleaded gasoline (the air). After the identification of leaded gasoline as a source of lead poisoning by US EPA, a major decline in their levels was found by replacing it with unleaded gasoline. Dongre [20] reported the toxicological profile, remedial solutions for lead levels in water by using polymeric materials, such as chitin and chitosan.
Zaltauskaite and Kniuipyte [21] reported the impact of lead concentration in soil on Eisenia fetida (earthworm). They found that lead in soil inhibits the growth of earthworms. Lead in soil can enter into human food by the vegetation in the contaminated soil [22]. The Types of carcinogenic effects of lead toxicity was explained in Figure 1. The lead toxicity in humans causes intestinal cancer, lung cancer, and central nervous system.
Figure 1.
Carcinogenic effect of Lead.
2.3 Sources, exposure and environmental impacts of cadmium
The major exposure of cadmium by human beings is through contaminated food and water. Cigarette smoke is the way of exposure through the inhalation route. The toxicity of cadmium is due to accumulation in plants and animals for nearly 25–30 years. Microbial fermentation is one of the effective methods to remove cadmium from food [23]. Another major source of cadmium in the environment is phosphate fertilizers and the waste incineration process. Blood cadmium levels are having a huge difference between smokers and non-smokers of cigarettes [1].
The presence of lead and cadmium in the human body can reach the brain and can cause Alzheimer’s disease [24]. After the exposure to the cadmium, in human, it can accumulate at the kidney, due to this reason urinary cadmium levels has been considered as biomarkers for cadmium levels in humans [25].
In case of occupational exposure to cadmium includes the workers at battery production, pigment industries, and electroplating. Because of long time accumulation in the human body even in small quantities is toxic and carcinogenic [26]. Another important source of cadmium in the soil is sewage sludge which can make the cadmium almost the same amount as fertilizers consumption (https://www.osti.gov/biblio/5478006, accessed on 1st October/2020). The types of carcinogenic effects of cadmium toxicity were explained in Figure 2. Cadmium shows its toxic effects on the gastric system and leads to gastric cancer, breast cancer, lung cancer, and it also affects the excretory system and leads to renal cancer.
Figure 2.
Carcinogenic effect of cadmium.
2.4 Sources, exposure, and environmental impacts of mercury
Mercury is the metal widely studied all over the world due to its toxic nature and easily entering into the food chain. An extensive review report was published by Jyothi and Mohamed Farook [6] regarding the sources, exposure, and toxicity of mercury. The toxicity of mercury depends on its chemical composition. Methyl mercury is more toxic than inorganic mercury. Due to its toxic nature and historical incidents like Minamata so many authors were published various facts regarding the sources, transport, and fate of mercury in the environment. Both volcanoes and forest fires are natural sources of mercury in the atmosphere. The burning of fossil fuels in power plants is one of the major anthropogenic sources of mercury (https://www.epa.gov/mercury/basic-information-about-mercury; accessed on 15th October/2020). Because of easy transportation, it is considered a global pollutant [27].
Even exposure to small quantities, shows toxic effects on various physiological systems, such as nervous and digestive systems and organs like lungs and kidneys. Due to this reason WHO declares mercury as one of the top most priority toxic metals (https://www.who.int/news-room/fact-sheets/detail/mercury-and-health; accessed on 20th October/2020). When it enters into water it largely affects the aquatic animal’s life and through them, it can enter into the food chain to reach human beings.
Yokoyama [28] reported the methylmercury poisoning control measures and the current situation of its effects on fetuses and infants particularly. In this study, they addressed the global cycle of methyl mercury also. Strode et al. [29] studied the emission of mercury into the North American atmosphere due to gold and silver mining in the 19th century. The types of carcinogenic effects of mercury toxicity was explained in Figure 3. Mercury toxicity effects on the rectal system and leads to colorectal cancer. It shows vast effects on the central nervous system leads to brain cancer and lung cancer.
Figure 3.
Carcinogenic effect of mercury.
2.5 Sources, exposure, and environmental impacts of arsenic
Arsenic is a metalloid but due to its toxic and carcinogenic nature, it is discussed under the heading of heavy metal toxicity. Abdul et al. [30] reviewed the health effects of arsenic exposure to human beings. According to this study, the majority of the population expose to this toxic metal through atmospheric air, groundwater, and certain kind of foods. The health effects are not limited to damage to cardiovascular, endocrine, renal, and reproductive systems. In various parts of the world such as India, Pakistan, and Bangladesh it was observed that major exposure to the arsenic is through groundwater. Shahid et al. [31] reported about the sources and health effects of arsenic through the exposure of groundwater in Pakistan. This study predicts nearly 47 million people of Pakistan at risk due to arsenic contamination in groundwater. They found that over 50% of the well they studied are having higher arsenic levels than WHO recommended levels in drinking water. A recent study [32] describes the occurrence and mobilization of arsenic in the groundwater of Bangladesh. In this study, they found that intensive usage of land for agriculture and usage of agrochemicals are the major reasons for arsenic contamination in groundwater of Bangladesh. Ahmed et al. [33] reported the situation of arsenic contamination in groundwater in a village in Bangladesh. For this purpose, they discussed 20 years situation of its exposure. Based on their results they found that the cancer risk to the population who are exposing to arsenic has 40% more than the non-exposures. The Types of carcinogenic effects of arsenic toxicity was explained in Figure 4. Sources and health effects of all the above discussed heavy metals are summarized in Table 2. Arsenic has its toxic carcinogenic effect on prostate glands and cause prostate cancer, leukemia and cause lesions in hepatic regions leads to cause of cancer of the liver.
Osteo related problems Prostate cancer Lung diseases Renal issues
Table 2.
Sources and health effects of heavy metals.
The permissible limits of lead, cadmium, arsenic, and mercury in different environmental matrices suggested by various international reputed agencies such as US EPA (Environmental Protection Agency), WHO (World Health Organization), and OSHA (Occupational Safety and Health Administration) are presented in Table 3.
US EPA
WHO
OSHA
Ambient Air
Drinking Water
Soil
Ambient Air
Drinking Water
Soil
Air at work place
Blood
Pb
0.151 μg/m3
151 μg/L
400 ppm1 (play areas); 1200 ppm non-play areas
—
152 μg/L
—
301 μg/m3
401 μg/dL
Cd
6.5–1306 ng /m3
0.0053 mg/L
854 mg/Kg
—
0.0033 mg/L
—
54 μg /m3
—
As
506 μg/m3
0.012 mg/L
—
—
0.012 mg/L
—
105 μg /m3
—
Hg
57 mg /m3
0.0022 mg/L
4–167 mg/Kg
—
0.0012 mg/L
—
—
—
Table 3.
Permissible limits of different toxic elements in environmental matrices.
The heavy metal toxicity and their environmental impact is a global issue due to their transportation through air, soil, and water. Based on various factors such as concentration and different major sources are the possible ways of entering the heavy metals through drinking water, air and foods. In minimum traces these metals are required for cellular, metabolic and hormonal functioning in humans but if the limitation exceeds its leads to the cause of severe hazardous effects in health. The toxicity of these metals is affecting the soil vastly by killing microorganisms present in soil which are very helpful to enhance fertility and nutrition levels of soils. According to the IARC, arsenic toxic effects are the cause of cancers in prostate glands, liver, blood, and skin. Mercury is the major reason for causing carcinogenic effects on the brain, lung, skin, and colorectal parts. The adverse effects of lead are the reason for intestinal, central nervous system, and lung cancers. The toxic effects of cadmium cause gastric, breast, lung, and renal cancers in humans.
Another diagnosis was identified in china, extreme high levels of lead toxicity in children were due to the pretty dish and in women, high lead levels in the blood is due to the usage of food storage cans. Cadmium is the major cause of Alzheimer disease and due to high usage of phosphate fertilizers they are accumulating in soils and entering into food chains. WHO states that mercury is hazardous toxic metal affecting aquatic life severely and consumption such mercury affected foods by human leads to several harmful diseases such as Minamata and cause several physiological effects.
\n',keywords:"heavy metals, heavy metal toxicity, sources, exposure, environmental impacts",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/74650.pdf",chapterXML:"https://mts.intechopen.com/source/xml/74650.xml",downloadPdfUrl:"/chapter/pdf-download/74650",previewPdfUrl:"/chapter/pdf-preview/74650",totalDownloads:95,totalViews:0,totalCrossrefCites:0,dateSubmitted:"July 20th 2020",dateReviewed:"December 4th 2020",datePrePublished:"December 30th 2020",datePublished:null,dateFinished:"December 30th 2020",readingETA:"0",abstract:"Heavy metals are defined in many ways, based on various factors such as density and atomic weight. Some of the heavy metals are essential as nutrients for humans such as iron, cobalt and, zinc in small quantities but are toxic in higher quantities. But few metals, such as lead, cadmium and, mercury are poisonous even in small quantities. The toxicity of heavy metals is depending on concentration,period of exposure and route of exposure. Heavy metal exposure takes place on human beings through inhalation from the atmosphere, intake through drinking water and, ingestion through the skin by dermal contact. The present chapter describes the definition of heavy metals, sources of these heavy metals, toxicity and, their impact on various environmental segments, such as air, water and, soil.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/74650",risUrl:"/chapter/ris/74650",signatures:"Narjala Rama Jyothi",book:{id:"10388",title:"Heavy Metals - Their Environmental Impacts and Mitigation",subtitle:null,fullTitle:"Heavy Metals - Their Environmental Impacts and Mitigation",slug:null,publishedDate:null,bookSignature:"Dr. Mazen Nazal",coverURL:"https://cdn.intechopen.com/books/images_new/10388.jpg",licenceType:"CC BY 3.0",editedByType:null,editors:[{id:"214815",title:"Dr.",name:"Mazen",middleName:null,surname:"Nazal",slug:"mazen-nazal",fullName:"Mazen Nazal"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}},authors:[{id:"303346",title:"Mrs.",name:"Narjala",middleName:null,surname:"Rama Jyothi",fullName:"Narjala Rama Jyothi",slug:"narjala-rama-jyothi",email:"ramadasaradhi@gmail.com",position:null,institution:null}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Types of heavy metals",level:"1"},{id:"sec_2_2",title:"2.1 Heavy metal toxicity",level:"2"},{id:"sec_3_2",title:"2.2 Sources, exposure, and environmental impacts of lead",level:"2"},{id:"sec_4_2",title:"2.3 Sources, exposure and environmental impacts of cadmium",level:"2"},{id:"sec_5_2",title:"2.4 Sources, exposure, and environmental impacts of mercury",level:"2"},{id:"sec_6_2",title:"2.5 Sources, exposure, and environmental impacts of arsenic",level:"2"},{id:"sec_8",title:"3. Conclusions",level:"1"}],chapterReferences:[{id:"B1",body:'Jarup, L. Hazards of heavy metal contamination. British Medical Bulletin 2003, 68(1), 167-182.'},{id:"B2",body:'Duffs, J.H. “Heavy metals” - a meaningless term? Pure and Applied Chemistry 2002, 74(5), 793-807.'},{id:"B3",body:'Emslay, J. Nature’s Building Blocks. Oxford university press, Oxford, 2011. ISBN: 978-0-19-960563-7.'},{id:"B4",body:'IARC. IARC monographs on the evaluation of carcinogenic risks to humans. Arsenic, metals, fibers and dusts. A review of human carcinogens. 2012, 100C.'},{id:"B5",body:'Lide, D.R. CRC Handbook of Chemistry and Physics. CRC Press, Florida. 2004. ISBN: 978-0-8493-0485-9.'},{id:"B6",body:'Jyothi, N.R., Mohamed Farook, N.A. Heavy metal toxicity in public health. 2020. IntechOpen, UK. DOI: http://dx.doi.org/10.5772/intechopen.90333.'},{id:"B7",body:'Suvarapu, L.N., Seo, Y.K., Baek, S.O. Heavy metals in the Indian atmosphere - a review. Research Journal of Chemistry and Environment. 2014, 18(8), 99-111.'},{id:"B8",body:'Suvarapu, L.N., Baek, S.O. Determination of heavy metals in ambient atmosphere : A review. Toxicology and Industrial Health 2017, 33(1), 79-96.'},{id:"B9",body:'Kim, J.J., Kim, Y.S., Kumar, V. Heavy metal toxicity : An update of chelating therapeutic strategies. Journal of Trace Elements in Medicine and Biology 2019, 54, 226-231.'},{id:"B10",body:'Giller, K., Witter, E., Mcgrath, S.P. Toxicity of heavy metals to microorganisms and microbial processes in agricultural soils: A review. Soil Biology and Biochemistry 1998, 30, 1389-1414.'},{id:"B11",body:'Yabe, J, Ishijuka, M., Umemura, T. Current levels in heavy metal pollution in Africa. Journal of Veterinary Medical Science 2010, 72, 1257-1263.'},{id:"B12",body:'Das, P., Samantaray, S., Rout, G.R. Studies on cadmium toxicity in plants : A review. Environmental Pollution 1997, 98, 29-36.'},{id:"B13",body:'Proshad, R., Kormoker, T., Mursheed, N., Islam, M.M., Bhuyan, M.I., Islam, M.S., Mithu, T.N. Heavy metal toxicity in agricultural soil due to rapid industrialization in Bangladesh: A review. International Journal of Advanced Geosciences 2018, 6(1), 83-88.'},{id:"B14",body:'Su, C., Jiang, L.Q., Zhang, W.J. A review on heavy metal contamination in the soil worldwide: Situation, impact and remediation techniques. Environmental Skeptics and Critics 2014, 3(2), 24-38.'},{id:"B15",body:'Bawa, U., Bukar, A., Abdullahi, Y. a review of lead poisoning, sources and adverse effects. ATBU journal of science, technology and Education 2015, 3(1), 71-79.'},{id:"B16",body:'Forsyth, J.E., Islam, M.S., Parvez, S.M., Raqib, R., Rahman, M.S., Muehe, E.M., Fendorf, S., Luby, S.P. Prevalence of elevated blood lead levels among pregnant women and sources of lead exposure in rural Bangladesh: A case control study. Environmental Research 2018, 166, 1-9.'},{id:"B17",body:'Ying, X.L., Gao, Z.Y., Yan, J., Zhang, M., Wang, J., Xu, J. Sources, symptoms and characteristics of childhood lead poisoning: Experience from lead specialty clinic in China. Clinical Toxicology 2018, 56(6), 397-403.'},{id:"B18",body:'Dong, C., Taylor, M.P., Gulson, B. A 25 year record of childhood blood lead exposure and its relationship to environmental sources. Environmental Research 2020, 186, 109357.'},{id:"B19",body:'Mahuta, M.A. An overlook of sources and strategies of minimizing lead poisoning in Nigeria. Journal of Humanities and Social Sciences 2020, 20(6), 330-340.'},{id:"B20",body:'Dongre, R.S. Lead: Toxicological profile, pollution aspects and remedial solutions. 2020. IntechOpen. DOI: 10.5772/intechopen.93095.'},{id:"B21",body:'Zaltauskaite, J., Kniuipyte, I., Kugelyte, R. Lead impact on the earthworm Eisenia fetida and earthworm recovery after exposure. Water, Air & Soil Pollution 2020, 49, 231.'},{id:"B22",body:'Sharma, N., Singh, A., Batra, N. Impact of soil, plant-microbe interaction in metal contaminated soils. Beneficial microbes for sustainable agriculture and Environmental Management, book chapter no.9. ISBN: 13: 978-1-77188-818-9.'},{id:"B23",body:'Genchi, G., Sinicropi, M.S., Lauria, G., Carocci, A., Catalano, A. The effects of cadmium toxicity. International Journal of Environmental Research and Public Health 2020, 17(11), 3782.'},{id:"B24",body:'Bakulski, K.M., Hu, H., Park, S.K. Chapter 51: Lead, cadmium and Alzheimer’s disease. The neuroscience of Dementia 2020, 2, 813-830.'},{id:"B25",body:'Suzzi, C.V., Kruse, D., Harrington, J., Levin, K., Meliker, J.R. is urinary cadmium a biomarker of long-term exposure in humans? A review. Current Environmental Health Reports 2016, 3, 450-458.'},{id:"B26",body:'Zhang, H., Reynolds, M. Cadmium exposure in living organisms: A short review. Science of the Total Environment 2019, 678, 761-767.'},{id:"B27",body:'Driscoll, C.T., Mason, R.P., Chan, H.M., Jacob, D.J., Pirrone, N. Mercury as global pollutant : Sources, pathways and effects. Environmental Science and Technology 2013, 47, 4967-4983.'},{id:"B28",body:'Yokoyama, H. Past, present and future of mercury pollution issues. In: Mercury Pollution in Minamata. Springer Briefs in Environmental Science. Springer, Singapore. 2018.'},{id:"B29",body:'Strode, S., Jaegle, L., Selin, N.E. Impact of mercury emissions from historic gold and silver mining: Global modeling. Atmospheric Environment 2009, 43(12), 2012-2017.'},{id:"B30",body:'Abdul, K.S.M., Jayasinghe, S.S., Chandana, E.P.S., Jayasumana, C., De Silva, P.M.C.S., Arsenic and health effects : A review. Environmental Toxicology and Pharmacology 2015, 40(3), 828-846.'},{id:"B31",body:'Shahid, M., Niazi, N.K., Dumat, C., Naidu, R., Khalid, S., Rahman, M.M., Bibi, I. A meta analysis of the distribution, sources and health risks of arsenic contaminated groundwater in Pakistan. Environmental Pollution 2018, 242, 307-319.'},{id:"B32",body:'Huq, M.E., Fahad, S., Shao, Z., Sarven, M.S., Khan, I.A., Alam, M., Saeed, M., Ulah, H. et. al. arsenic in groundwater environment in Bangladesh : Occurrence and mobilization. Journal of Environmental Management 2020, 262, 110318.'},{id:"B33",body:'Ahmed, S.A., Faruquee, M.H., Khan, M.H. Twenty years of arsenic contamination and Arsenicosis patients in a village of Bangladesh. American Journal of Public Health Research 2020, 8(6), 190-196.'},{id:"B34",body:'Ebrahimi, S. J. A., Eslami, A., Ebrahimjadeh, L. Evaluation of heavy metals concentration in the drinking water distribution network in Kurdistan villages in the year 2012. Research journal of pharmaceutical, biological and chemical Sciences 2015, 6(2), 55-61.'},{id:"B35",body:'Ye, B.J., Kim, B.G., Jeon, M.J., Kim, S.Y., Kim, H.C., Jang, T.W., Chae, H.J., Choi, W.J., Ha, M.N., Hong, Y.S. Evaluation of mercury exposure level, clinical diagnosis and treatment for mercury intoxication. Annals of Occupational and Environmental Medicine 2016, 28, 5.'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Narjala Rama Jyothi",address:"ramadasaradhi@gmail.com",affiliation:'
Department of Basic Science and Humanities, Sri Padmavathi Mahila Visvavidyalayam, Tirupati, Andhra Pradesh - 517501, India
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He received his Ph.D. from Chonbuk National University, South Korea, in 2007. After obtaining his Ph.D., Dr. Rahman worked as a postdoctoral fellow and assistant professor in pioneer research centers and universities in South Korea, Japan, and Saudi Arabia (2007–2011). Presently, he is a full professor at the Center of Excellence for Advanced Materials Research (CEAMR) and the Chemistry Department at King Abdulaziz University, Jeddah, Saudi Arabia. He has published many international and domestic articles and book chapters as well as edited several books. 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Highlights and importance are laid on real or practical problems, ranging from chemical application to biomedical monitoring, from in vitro to in vivo, and from single cell to animal to human measurement. This offers a unique opportunity of exchanging and combining the scientist or researcher in electrochemical sensors in largely chemistry, biological engineering, electronic engineering, and biomedical and physiological fields.",editors:[{id:"24438",title:"Prof.",name:"Mohammed Muzibur",surname:"Rahman",slug:"mohammed-muzibur-rahman",fullName:"Mohammed Muzibur Rahman"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",title:"Edited Volume"}},{type:"book",slug:"advances-in-colloid-science",title:"Advances in Colloid Science",subtitle:null,coverURL:"https://cdn.intechopen.com/books/images_new/5403.jpg",abstract:"This book Advances in Colloid Science covers a number of up-to-date research advancement and progresses on colloids. 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The innovative frontiers are branching out from time to time to advanced nanotechnology. It is an important booklet for scientific organizations, governmental research-centers, academic libraries, and the overall research and development of nano-materials in general. It has been created for widespread audience with diverse backgrounds and education.',editors:[{id:"24438",title:"Prof.",name:"Mohammed Muzibur",surname:"Rahman",slug:"mohammed-muzibur-rahman",fullName:"Mohammed Muzibur Rahman"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",title:"Edited Volume"}}],chaptersAuthored:[{title:"Iron Oxide Nanoparticles",slug:"iron-oxide-nanoparticles",abstract:null,signatures:"Mohammed M. Rahman, Sher Bahadar Khan, Aslam Jamal, Mohd Faisal and Abdullah M. 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His research\nfocuses on photonic materials including metamaterials, quantum\ndots, and plasmonic nanomaterials that can be used in a wide\nrange of nanophotonics applications. His recent interests also\ninclude nano-bioimaging, 3D printing, nanostructures for tissue\nengineering (ZnO, TiO2, etc.) and biomaterials including carbon, graphene, and\ndiamond quantum dots. He is an editorial board member and reviewer for different\ninternational journals and has collaborated with local and international academics/\nresearchers on post-graduate research projects. He has edited four books and published four chapters and more than 105 articles in scientific journals and reviewed\nconference proceedings. His papers have been cited more than 2100 times with\nh-index 23 and i-10 index 41 (Google Scholar).",institutionString:"Nuclear Science and Technology Research Institute",institution:null},{id:"69408",title:"Prof.",name:"Raphael",surname:"Schneider",slug:"raphael-schneider",fullName:"Raphael Schneider",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Laboratoire Réactions et Génie des Procédés",institutionURL:null,country:{name:"France"}}},{id:"94041",title:"MSc.",name:"Nicoleta",surname:"Petrea",slug:"nicoleta-petrea",fullName:"Nicoleta Petrea",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"CBRNE (United Kingdom)",institutionURL:null,country:{name:"United Kingdom"}}},{id:"102433",title:"Mr.",name:"Hamed",surname:"Mehranpour",slug:"hamed-mehranpour",fullName:"Hamed Mehranpour",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"118068",title:"Dr.",name:"Aslam",surname:"Jamal",slug:"aslam-jamal",fullName:"Aslam Jamal",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Najran University",institutionURL:null,country:{name:"Saudi Arabia"}}},{id:"118069",title:"Dr.",name:"Sher",surname:"Khan",slug:"sher-khan",fullName:"Sher Khan",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Najran University",institutionURL:null,country:{name:"Saudi Arabia"}}},{id:"118070",title:"Dr.",name:"M",surname:"Faisal",slug:"m-faisal",fullName:"M Faisal",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Najran University",institutionURL:null,country:{name:"Saudi Arabia"}}},{id:"118319",title:"MSc.",name:"Rodica Mihaela",surname:"Lungu",slug:"rodica-mihaela-lungu",fullName:"Rodica Mihaela Lungu",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"CBRNE (United Kingdom)",institutionURL:null,country:{name:"United Kingdom"}}},{id:"136004",title:"Dr.",name:"Masoud",surname:"Askari",slug:"masoud-askari",fullName:"Masoud Askari",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null}]},generic:{page:{slug:"OA-publishing-fees",title:"Open Access Publishing Fees",intro:"
The Open Access model is applied to all of our publications and is designed to eliminate subscriptions and pay-per-view fees. This approach ensures free, immediate access to full text versions of your research.
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XML Typesetting and pagination - web (PDF, HTML) and print files preparation
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Permanent and unrestricted online access to your work
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For Authors who are still unable to obtain funding from their institutions or research funding bodies for individual projects, IntechOpen does offer the possibility of applying for a Waiver to offset some or all processing feed. Details regarding our Waiver Policy can be found here.
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Proven world leader in Open Access book publishing with over 10 years experience
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The Open Access Publishing Fee (OAPF) is payable only after your full chapter, monograph or Compacts monograph is accepted for publication.
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1,400 GBP Chapter - Edited Volume
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10,000 GBP Monograph - Long Form
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4,000 GBP Compacts Monograph - Short Form
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*These prices do not include Value-Added Tax (VAT). Residents of European Union countries need to add VAT based on the specific rate in their country of residence. Institutions and companies registered as VAT taxable entities in their own EU member state will not pay VAT as long as provision of the VAT registration number is made during the application process. This is made possible by the EU reverse charge method.
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Services included are:
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An online manuscript tracking system to facilitate your work
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Personal contact and support throughout the publishing process from your dedicated Author Service Manager
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Assurance that your manuscript meets the highest publishing standards
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English language copyediting and proofreading, including the correction of grammatical, spelling, and other common errors
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XML Typesetting and pagination - web (PDF, HTML) and print files preparation
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Discoverability - electronic citation and linking via DOI
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Permanent and unrestricted online access to your work
What isn't covered by the Open Access Publishing Fee?
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If your manuscript:
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\n\t
Exceeds 20 pages (for chapters in Edited Volumes), an additional fee of 40 GBP per page will be required
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If a manuscript requires Heavy Editing or Language Polishing, this will incur additional fees.
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Your Author Service Manager will inform you of any items not covered by the OAPF and provide exact information regarding those additional costs before proceeding.
\n\n
Open Access Funding
\n\n
To explore funding opportunities and learn more about how you can finance your IntechOpen publication, go to our Open Access Funding page. IntechOpen offers expert assistance to all of its Authors. We can support you in approaching funding bodies and institutions in relation to publishing fees by providing information about compliance with the Open Access policies of your funder or institution. We can also assist with communicating the benefits of Open Access in order to support and strengthen your funding request and provide personal guidance through your application process. You can contact us at oapf@intechopen.com for further details or assistance.
\n\n
For Authors who are still unable to obtain funding from their institutions or research funding bodies for individual projects, IntechOpen does offer the possibility of applying for a Waiver to offset some or all processing feed. Details regarding our Waiver Policy can be found here.
\n\n
Added Value of Publishing with IntechOpen
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Choosing to publish with IntechOpen ensures the following benefits:
\n\n
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Indexing and listing across major repositories, see details ...
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Long-term archiving
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Visibility on the world's strongest OA platform
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Live Performance Metrics to track readership and the impact of your chapter
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Dissemination and Promotion
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Benefits of Publishing with IntechOpen
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Proven world leader in Open Access book publishing with over 10 years experience
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Most competitive prices in the market
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Optimized processes, enabling publication between 8 and 12 months
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Personal support during every step of the publication process
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+146,150 citations in Web of Science databases
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Currently strongest OA platform with over 150 million downloads
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