SVC flow velocities changes at different stages of SVCO (
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More than half of the publishers listed alongside IntechOpen (18 out of 30) are Social Science and Humanities publishers. IntechOpen is an exception to this as a leader in not only Open Access content but Open Access content across all scientific disciplines, including Physical Sciences, Engineering and Technology, Health Sciences, Life Science, and Social Sciences and Humanities.
\\n\\nOur breakdown of titles published demonstrates this with 47% PET, 31% HS, 18% LS, and 4% SSH books published.
\\n\\n“Even though ItechOpen has shown the potential of sci-tech books using an OA approach,” other publishers “have shown little interest in OA books.”
\\n\\nAdditionally, each book published by IntechOpen contains original content and research findings.
\\n\\nWe are honored to be among such prestigious publishers and we hope to continue to spearhead that growth in our quest to promote Open Access as a true pioneer in OA book publishing.
\\n\\n\\n\\n
\\n"}]',published:!0,mainMedia:null},components:[{type:"htmlEditorComponent",content:'
Simba Information has released its Open Access Book Publishing 2020 - 2024 report and has again identified IntechOpen as the world’s largest Open Access book publisher by title count.
\n\nSimba Information is a leading provider for market intelligence and forecasts in the media and publishing industry. The report, published every year, provides an overview and financial outlook for the global professional e-book publishing market.
\n\nIntechOpen, De Gruyter, and Frontiers are the largest OA book publishers by title count, with IntechOpen coming in at first place with 5,101 OA books published, a good 1,782 titles ahead of the nearest competitor.
\n\nSince the first Open Access Book Publishing report published in 2016, IntechOpen has held the top stop each year.
\n\n\n\nMore than half of the publishers listed alongside IntechOpen (18 out of 30) are Social Science and Humanities publishers. IntechOpen is an exception to this as a leader in not only Open Access content but Open Access content across all scientific disciplines, including Physical Sciences, Engineering and Technology, Health Sciences, Life Science, and Social Sciences and Humanities.
\n\nOur breakdown of titles published demonstrates this with 47% PET, 31% HS, 18% LS, and 4% SSH books published.
\n\n“Even though ItechOpen has shown the potential of sci-tech books using an OA approach,” other publishers “have shown little interest in OA books.”
\n\nAdditionally, each book published by IntechOpen contains original content and research findings.
\n\nWe are honored to be among such prestigious publishers and we hope to continue to spearhead that growth in our quest to promote Open Access as a true pioneer in OA book publishing.
\n\n\n\n
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Two major topics were covered in this chapter. The first was to explore a method of establishing a rabbit model of SVCS by injecting VX2 tumor cell suspension transcutaneously under ultrasound guidance, and to observe the radiotherapeutic effects by SVC Doppler flow changes. The establishment of this model would offer an experimental evidence for better diagnosis, treatment and follow-up observation of SVCS. The second was to investigate the evolution of the SVC Doppler flow changes in patients with SVCS and its value in assessing clinical therapeutic effects in these patients.
An animal SVC obstruction model would be of use in further studies directed at optimizing the therapeutic strategies for original tumors causing SVC syndrome in patients, yet no detailed information about the establishment of an animal SVC obstruction (SVCO) model can be found in the literature. It has been reported that VX2 tumor cell could be successfully inoculated into multiple organs to induce malignant tumors in rabbits (De Crespigny et al, 1999; Goldberg et al, 1999; Ishida et al, 2000), and various imaging modalities has been adopted to assess the effect of this inoculation (Kim et al, 2000; Liu et al, 2001). We hypothesize that a rabbit model with SVCO could be established by inoculating VX2 tumors cells into the areas around the SVC.
Superior vena cava syndrome (SVCS) is a clinical expression of obstruction of blood flow through the SVC, and more than 80% cases are caused by malignant tumors. Though contrast-enhanced spiral or multi-slice CT is now able to identify accurately the site of occlusion or stenosis, it has been greatly limited by its high cost and radiation and thus is not appropriate for follow-up observations. In contrast, ultrasound is safe, reproducible, and relatively inexpensive. Doppler ultrasonography has been successfully used to assess the rabbit SVC obstruction model. We hypothesize that Doppler flow patterns of SVC could be applied for assessing the severity of SVCS and its therapeutic effects.
There are three objectives for the animal study. The first is to study the feasibility of establishing a model of SVCO in rabbits by infusing VX2 tumor cell suspension transcutaneously with ultrasound guidance, and to evaluate the applications of this animal model. The second is to study morphologic and hemodynamic changes of superior vena cava (SVC) in rabbits with SVCO using two-dimensional and Doppler ultrasound and to explore the relationship between the tumor size in specimen and the two-dimensional and Doppler ultrasonographic characteristics. The third is to analyze the ultrasonographic characteristics and histopathological changes after radiotherapy in rabbits with SVCO, and to provide useful information for assessing the SVC syndrome therapeutic effect in clinic.
The objective of the human study was to evaluate the Doppler SVC’s flow patterns and their value in assessing SVCS.
Fifteen adult healthy New Zealand White rabbits were enrolled in this study. Anesthesia was performed by injecting Ketamine Hydrochloride into the posterior leg muscles of rabbits at a dose of 30mg/kg. The VX2 tumor cell suspension was prepared under the sterile condition. About 0.1ml tumor cell suspension was infused transcutaneously in front of SVC and close to its anterior wall guided by ultrasound. The SVC morphology and hemodynamics as well as the tissues around SVC were examined with two-dimensional and Doppler ultrasonography once every 3 days from the 9th day after the injection of tumor cell suspension till the natural death of the rabbits. These findings were compared with those by CT and digital subtraction angiography (DSA).
One rabbit did not develop tumor after the injection of VX2 tumor cell suspension and thus excluded. The rest of the fourteen rabbit models of SVCO were examined by two-dimensional and Doppler ultrasonography using Sequoia 512 computed ultrasonograph with the probe of 7v3c.
The sizes of the tumor and the SVC morphology and hemodynamics were observed with the transducer placed in the right supraclavicular region view using Sequoia 512 computed ultrasonograph once every 3 days from the 12th day after the injection, and compared with the ultrasonographic findings before injection. The ultrasonographic findings were also compared with those of CT and autopsy findings, respectively.
Thirteen survived rabbits with SVCO due to VX2 tumor were enrolled in this study. The tumors in mediastina were exposed one minute with 2gy everyday by SIEMNS MEVATRON 6745, which was 10 times in all. Before the radiotherapy and On the 10, 17 and 24 day after radiotherapy, the sizes of the tumor and SVC, SVC velocities and echo characteristics in the 13 rabbits were detected with the transducer placed in the right supraclavicular region using Sequoia 512 computed ultrasonograph. The mediastina neoplasm tissue before and after radiotherapy were sampled for HE staining and TUNEL for analysis of the number of the apoptotic cells.
Forty-two patients (26 females and 16 males) with SVCS caused by neoplasm (34 with right upper lung cancer and 8 with mediastinal tumor), aged from 22 to 56 years old were included in this study when they were scheduled for radiotherapy and/or chemotherapy treatment from January 2000 to June 2005. All the patients had upper extremity and facial swelling at initial diagnosis. Twenty volunteers (15 males and 5 females) aged 23–52 years with no history of cardiac and pulmonary diseases were recruited as controls. All informed consents of the patients were acquired.
Acuson Sequoia 512 ultrasonograph equipped with7V3C transducer was used. Electrocardiogram and respiratory curve were recorded simultaneously. The following procedures were in accordance with the ethical standards of the committee on human experimentation of the institution and approved by Tangdu Hospital committee.
Right supraclavicular approach. Patients took a supine position. With the transducer placed in the fossa between the sterna and clavicular heads of the sternomastoid muscle, the upper part of the SVC and its adjacent structures were fully displayed.
Subcostal approach. Patients took a supine position. With the transducer placed in the subcostal region, the lower part of SVC was displayed.
The SVC spectra were recorded. The flow velocities of the two forward waves, systolic wave (S) and diastolic wave (D) and the two reversed waves during ventricular and atrial contraction (VR and AR), were measured. The variation of these flow velocities with cardiac cycle and respiration was analyzed with SPSS software. All the patients were followed up for more than 11 months.
Fourteen rabbits were found to have the tumors para-SVC and/or in the SVC cavity by ultrasonography. One rabbit dropped off because no tumor grew until 42nd day after the infusion. The success rate of developing rabbit SVCO model was about 93.33%.
The diameters of the tumors were (80.70±4.28) mm. With the tumor growing, the lumen of SVC was deformed and narrowed, and the wall of SVC was disrupted shown by two-dimensional ultrasonography (Figure 1 and 2). The tumor size growth was linearly correlated with time, and the correlation coefficient was 0.9855 (Figure 3). The tumor diameter by ultrasound was similar to the diameter by autopsy ((80.70±4.28mm vs. 82.16±3.41mm, t=0.998, P=0.327405).
Two-dimensional ultrasonography of normal SVC
Two-dimensional ultrasonography of a rabbit with SVCO
Tumor growing with time
All the 14 rabbits developed SVCO. In eight rabbits, SVC was found to be oppressed by two-dimensional ultrasonography on the 15th day after infusion, Aliasing mosaic flow signals and high flow velocity spectra in SVC were demonstrated by Doppler ultrasonography. In another 4 rabbits, the stenotic blood flow velocities were displayed on the 18th day after infusion. In the last 2 rabbits, the abnormal blood flow was seen on the 21st and the 24th day, respectively.
The development of SVCO could be divided into three stages. Early stage: About 1 week after. The tumor area was 0.5 cm2~3cm2. The echotexture was hpyoechoic and evenly distributed, the shape of the tumor was regular with pseudocapsule. SVC was compressed, but the SVC wall was relatively intact (Figure 4). Mid stage: the tumor area was 3.1 cm2~6.0 cm2. Most of the tumors still were hypoechoic echotexture. Hyperechoic textures could be seen within some tumors. T shape of the tumor was not regular. At this stage, the SVC was oppressed and its course became abnormal; the lumen of SVC was narrowed, and the wall was infiltrated by tumors (Figure 5). Late stage: The tumor area reached more than 6.1 cm2, and the shape was dramatically irregular. Mixing echotexture was seen, but was mainly hypoechoic. SVC was severely oppressed and correlated well with the size of the tumor (Figure 6). The ultrasonographic findings of SVCO at the late stage correlated well with those findings by CT and the autopsy.
SVC being compressed by a mass (M) in a rabbit with SVCO. The wall of SVC is intact
SVC being compressed in a rabbit with SVCO. The wall of SVC is infiltrated by the mass (M)
Relationship between the tumor size and the SVC diameter
In SVCO, the color Doppler ultrasound showed mosaic or weak or even no flow signals within SVC (Figure 7). In normal rabbits, the pulsed Doppler ultrasound showed laminar flow in SVC (Figure 8); while in rabbits with SVCO, the SVC flow was turbulent and the spectral window was disappeared (Figure 9). The peak flow velocities of SVC waves were less influenced by respiratory cycle in SVCO compared to normal (Figure 10). SVC flow velocities significantly increased during early and mid stages of SVCO (Table 1).
S | D | VR | AR | |
Early and middle stages | 78.25±14.97 | 59.68±13.16 | 19.22±4.99 | 17.44±2.67 |
Late stage | 33.71±18.90 | 33.55±20.03 | 10.53±3.27 | 9.33±1.58 |
T | 6.912 | 4.079 | 5.450 | 9.781 |
P | <0.0001 | 0.0004 | <0.0001 | <0.0001 |
SVC flow velocities changes at different stages of SVCO (
Mosaic and thread-like color Doppler flow signals were seen in SVC in a rabbit with SVCO by color Doppler ultrasonography. T: tumor
Normal SVC Doppler flow
Increased SVC Flow velocities in the same rabbit as shown in figure 7
Peak velocities of SVC waves were less influenced by respiratory cycle than normal
The tumor showed a tendency to get smaller after radiotherapy. The echotexture of the tumor partly turned to be hyperechoic. The diameter of SVC become bigger and the flow velocities decreased at the site of tumor compared with that before the radiotherapy (Figure 11 and 12). The tumor diameter, SVC diameter and velocity changes after radiotherapy were shown in table 2.
SVC Doppler flow waveforms before the radiotherapy
SVC Doppler flow velocities decreased after radiotherapy from the same patient shown in figure 11
D1 (mm) | D2(mm) | SysVmax(cm/s) | |
Before radiotherapy | 17.86±1.12 | 2.65±0.32 | 87.38±12.94 |
10d after radiotherapy | 17.64±1.08 | 2.78±0.37 | 84.65±11.46 |
17d after radiotherapy | 17.13±1.18 | 2.94±0.93 | 81.37±11.50 |
24d after radiotherapy | 16.38±1.60 | 3.23±0.28 | 77.55±12.34 |
Tumor diameter (D1), SVC diameter (D2) and systolic maximal flow velocity (SysVmax) changes after radiotherapy
HE staining and TUNEL assay showed that the number of apoptotic cells in the tumor was much more than that before the radiotherapy (Figure 13& 14) (P<0.01).
HE staining showing that the apoptotic cells increased after radiotherapy (Right) compared to those before (Left)
TUNEL assay showing that the apoptotic cells increased after radiotherapy (Right) compared to those before (Left)
Different from healthy subjects, where laminar flow was demonstrated (Figure 15), the SVC flow spectra in patients with mild SVCS showed turbulent flow (Figure 16) and the spectral window was disappeared. In patients with moderate degree of SVCS, the distinct biphasic forward waves (S- and D-waves) of SVC were lost (Figure 17). In addition, we found that the smaller of the VR- and AR-waves were, the farther the oppressed segment of SVC was away from the right atrium.
SVC Doppler flow spectra in healthy subjects. Doppler interrogation of the SVC shows systolic and diastolic phases of flow (S- and D-waves) toward the heart and late ventricular systolic and atrial systolic phases of backward flow (VR- and AR-waves)
SVC Doppler flow spectra in mild SVCS. The forward waves showed high velocity and no distinct spectral window
SVC Doppler flow spectra in severe SVCS. The S, D AR, and VR waves could not be identified
The RVI in SVCS group were much lower than those in the control group (S-wave: 1.9±3.6 %vs.16.34±8.96%, P = 0.0003; D-wave: 2.80±1.23% vs. 26.32±42%, P =0.0087). After treatment, the flow velocities of SVC decreased significantly with each month and the RVI was significantly increased compared to those before the treatment.
The animal study demonstrated that SVCO rabbit model could be easily established transcutaneously with the guidance of ultrasound. With VX2 tumor growing, SVC was oppressed and/or infiltrated by the tumor tissues and ultimately resulted in the blockage of the SVC, and developed SVCO. The pathogenesis of SVCO in rabbits was similar to that of SVCO in humans, indicating that this rabbit model of SVCO caused by VX2 tumor could be an ideal model of SVCO.
Two-dimensional ultrasonography could be used to demonstrate the morphologic changes of the tumor and SVC. Color Doppler could sensitively detect the abnormal blood flow of SVC in rabbits with SVCO. It could be mosaic, weak or even no flow signals depending on the different stages and severity of SVCO. Doppler flow spectra of SVC showed less respiratory influences on the flow velocities in rabbits with SVCO.
Linear accelerator radiotherapy could lead to cell apoptosis of the VX2 tumor transplanted besides SVC. HE staining and TUNEL assay showed increased number of apoptotic cells of the tumor tissue compared to that before radiotherapy. With the apoptosis of the tumor cells, the echotexture of the tumor partly turned to be hyperechoic and the tumor itself was getting smaller. Thereby the oppression severity of the SVC was decreased, the diameter of SVC was getting bigger, and finally the obstruction was getting resolved.
With the release of the obstruction after radiotherapy, the respiratory variation of the SVC Doppler flow velocities could be recovered and the peak flow velocities of SVC decreased gradually after the treatment shown by Doppler ultrasonography. However, the tumor size, SVC diameter and the flow velocity could not completely get back to normal after treatment. This suggests that Linear accelerator radiotherapy could not inactivate VX2 tumor completely, which might be related to the high grade of malignancy, poor proliferation of the VX2 tumors.
The human study demonstrates that respiratory variations of SVC Doppler flow velocities are significantly decreased in patients with SVCS, but could be recovered gradually after treatment, indicating that respiratory variations of Doppler flow changes of SVC correlate well with the severity of SVCS, and may be used to assess the therapeutic effects of SVCS.
The SVC Doppler flow patterns in patients with SVCS were characterized with fill-in and broaden spectra, nondistinct outlines. The D-wave was unable to return to the baseline at end diastole, which was consistent with previous studies (Yano&Shimada, 1997). When the patients’ condition improved after treatment,, the pressure gradient at the stenotic segment of SVC also decreased, indicating that SVC hemodynamic changes was in accordance with the clinical courses of SVCS (Behar et al., 2001; Tighe et al., 2000).
It has been well known that, respiration had significant influence on SVC flow velocities in healthy subjects. The S- and D-wave peak velocities are much greater on inspiration compared with those on expiration because the decreased intrathoracic pressures during inspiration causes increased venous return (Jellinek et al., 2000; Vieillard et al., 2001). In patients with SVCS, However, the obstruction prevents the conduction of the intrathoracic pressure changes into the right atrium and to SVC, resulting in decreased or even diminished respiratory variations of S- and D-wave flow velocities. Once this obstruction is released after treatment, the SVC Doppler flow velocities would decrease and the respiratory variations of these flow velocities would be back.
Rabbit model of SVCO caused by VX2 tumor could be an ideal model of SVCO for clinical study. Doppler ultrasound is the first method of choice for assessing the hemodynamics of SVCO.
Normally, respiration had significant influence on SVC morphology and dynamics in healthy subjects. This influence would become less in the patients with SVCS, suggesting that SVC Doppler spectra could reflect the severity of SVCS.
Increase in global temperature had major impact on crop productivity especially in tropical and sub tropical regimes. Based on climate model predictions, around 1.8–4.0°C rise in air temperature was expected in 21st century [1]. The increase in temperature beyond a certain threshold level tends to induce detrimental effects in plant growth and development. In general, the elevation in temperature of 10–15°C above ambient triggers heat shock in crop plants. The extent of induced heat stress depends on the duration, intensity and rate of increase in global air temperature [2]. Indian lowlands share 15 per cent of global wheat production. The change in global climate would shift these fertile lowlands into heat stressed unproductive environment [3]. Similarly, the cultivation of cereals in Southern Africa and South East Asia was predicted to be heat stressed zone in near future [4]. Around 4–14% yield decline in rice was encountered due to elevated temperature of 1°C in South-East Asia [5]. The declined productivity due to elevated temperature imposes the urgent need for development of climate resilience genotypes. Evolving heat tolerant cultivars would highly benefit the livelihood of developing countries as around 70–80% of population relies on agriculture. Understanding the effect of heat stress on crop plants and its adaptation mechanisms would help in framing out the breeding strategies for high temperature tolerance.
\nHeat tolerance in crop plants is a complex mechanism involving adaptations through altered physiological process, morpho-anatomical features and induction of several biochemical pathways. On exposure to high temperature, several signal transduction pathways were triggered leading to changes in gene expression. As a result, varied stress related proteins were synthesized contributing heat tolerance in plants [6]. The tolerance mechanism to high temperature stress varies within genotypes of a plant species. The existing variation between and within species provide scope for evolving heat tolerant lines through conventional breeding approaches [7]. Dissecting out genetic information through molecular tools would hasten the development of climate resilient cultivars contributing to food security in near future. A brief review on plant response, adaptation mechanisms and genetic approaches to combat heat stress were presented in this chapter.
\nHeat stress had varying impact on different phenological stages viz., germination, seedling, vegetative, flowering and reproductive of crop plants [8]. The plant response to heat stress depends on the duration, degree of rise in temperature and plant type. Under tropical regimes, high temperature with intense solar radiation poses a major limiting factor for yield by inducing leaf abscission, leaf senescence, scorching of leaves, branches and stems, growth inhibition, pollen infertility and poor seed formation [9, 10]. A significant decline in relative growth rate, shoot dry weight and net assimilation rate was recorded in sugarcane, maize and pearl millet on exposure to high temperature stress [11]. High reduction in grain quality was recorded in most of the cereal crops grown under heat stress environments [12]. Several physiological processes such as partitioning of assimilates, plant-water relations and shoot growth was affected due to heat stress in common bean [13]. In general, the susceptibility to heat stress was found higher at reproductive stage of plant development. An excessive yield loss is recorded in legumes on exposure to high temperature (30–35°C) during anthesis stage [14]. Drastic reduction in grain number and weight was observed in wheat at high temperature regimes [15]. Heat stress affects several metabolic pathways leading to accumulation of reactive oxygen species (ROS) which is a major component for oxidative stress in crop plants [16]. The photosystem centres (PS I and PS II) of chloroplast, mitochondria and peroxisomes are the major sites for generation of ROS in plants [17]. High temperature stress disrupts the stability of cell membrane through protein denaturation [18]. The induction of ROS due to high temperature stress was correlated with premature leaf senescence in Gossypium sp. [19]. Accumulation of ROS in root cells was evidenced in wheat on exposure to high temperature for two days [20].
\nPlants tend to adapt several complex mechanisms through phenological and morphological changes to combat high temperature stress (Figure 1). On heat stress regimes, plants exhibit varied short term escape/avoidance mechanisms viz., altered leaf orientation, transpirational cooling, altered membrane lipid properties, early maturation and so on for its survival. Plants show varied degree of leaf rolling upon intensity of solar radiation. A significant tolerance to high temperature was observed in wheat by maintenance of water potential in flag leaf through adoption of leaf rolling under heat shock conditions [21]. Increase in trichomatous and stomatal densities, waxy layer on leaves, and larger xylem vessels are the common features induced during heat stress [22]. On contrary, plants also evolve long term tolerance mechanisms for its effective survival and productivity under high temperature. Induction of osmoprotectants, antioxidants, late embryogenesis abundant proteins, dehydrins, and heat shock proteins are the major factors involved in counteracting the heat shocks. Accumulation of osmolytes such as proline, trehalose, and glycine betaine plays a vital role in imparting tolerance via cellular osmotic adjustment, detoxification of ROS, stabilization of enzymes and membrane proteins [23]. Several enzymatic and non-enzymatic antioxidant defense components are also involved in protection against oxidative stress induced by free radicals [24]. The activities of ROS scavenging enzymes are temperature specific. In general, most of the antioxidant enzymes show increased activity with elevation in temperatures. It is also influenced by genotype, growing season and phenological stages of plant [25]. Under high temperature conditions, several signaling molecules such as nitrous oxide, Ca-dependent protein kinases, Mitogen mediated protein kinase, sugars, and phytohormones play a role in stimulation of stress responsive genes via transduction pathways [26]. Evolving adaptation mechanisms (either tolerance or avoidance) to high temperature and drought would be more rewarding at arid conditions as it is often correlated.
\nAdaptation mechanisms for high temperature tolerance in crop plants.
Breeding for high temperature tolerance requires an essential knowledge on plant adaptation response to heat shocks. In general, the genotypes exhibiting less detrimental effect on photosynthesis and reproductive development tend to survive well under heat prone areas [27]. Involvement of these two components in selection criteria would be beneficial in evolving thermo tolerant cultivars. Tolerant genotypes evolve several morphological, physiological and biochemical alterations in response to heat shocks. Knowledge on sensitivity of several phenological stages to high temperature will pave way for trait specific improvement. High temperature is often correlated with other environmental factors which poses a major limitation for selection under field conditions. At present, varied selection criteria has been developed by scientists, which favors delineation of superior variety at prevailing environment [28]. Heat tolerant index has been evolved for sorghum which depicts the proportion of growth recovery after exposure to high temperature stress. It is the ratio of increase in coleoptile growth in a heat stress environment [50°C] to the enhancement in coleoptile length under normal environment (non-stress) [29]. It proves cost effective and rapid method to screen a large population size within shorter period. A proper validation of such technique would facilitate the development of tolerant lines in other crop species. Pollen viability and fruit set was considered as major selection criteria to predict yield under high temperature stress in tomato [30]. Physiological based trait selection such as harvest index, photosynthetic efficiency, respiration rate, delayed senescence and canopy architecture will also contribute towards increased tolerance to heat stress [31, 32].
\nInter-mating among closely related individuals for improvement of economic traits resulted in decline of genetic variability in a crop species [33]. Characterization of gene pool including land races and wild relatives would offer several tolerant genes for abiotic tolerance. Extensive efforts were made in screening of heat tolerant genotypes which can be directly introduced as a cultivar or utilized to introgress gene into new genetic background [34]. Thermo-tolerant lines were successfully isolated from wild gene pool in wheat [35]. High magnitude of variation was observed in wild progenitor “Aegilops tauschii” of wheat for cell viability and membrane stability [36]. Similarly, a heat tolerant source for reproductive stage was identified in A. geniculata and A. speltoides Tausch which would pave way in development of thermo-tolerant hexaploid wheat cultivars in near future [37]. A higher growth rate and improved photosynthetic efficiency was observed in wild relative “Oryza meridionalis” of rice at high temperature [38]. Indirect selection on pollen viability led to identification of thermo-tolerant accessions in soybean (DG 5630RR) [39], chickpea (ICC15614 & ICC1205) [40], maize (AZ100) [41], and several other crop species. Direct selection based on yield under target environment (heat stress) resulted in development of tolerant lines in many tropical grain legumes. Four tolerant genotypes/accessions viz., SRC-1-12-1-48, SRC-1-12-1-182, 98012-3-1-2-1 and 98020-3-1-7-2 were isolated in common bean by employing stress tolerant indices [42]. Nine thermo-tolerant wild accessions were delineated in USDA upland cotton germplasm by employing chlorophyll fluorescence technique [43].
\nEvolving thermo-tolerance through conventional breeding approach proves promising in many crop species. Breeding for early maturing genotype in broccoli had improved head quality by avoiding heat stress at flowering stage [44]. In general, breeding programmes are carried out in hotter regions which promote selection of thermo-tolerant traits. Physiological based trait breeding was practiced at International Maize and Wheat Improvement Center (CIMMYT) for development of heat tolerant cultivars in wheat. The parental genotypes were characterized through various crossing schemes and appropriate breeding programme was framed for improvement of thermo related traits [45]. A wild ancestor “T. tauschii” was utilized as a gene donor for achieving increased grain size and filling percent under high temperature through recurrent selection [46]. Similarly, three cycles of recurrent selection had led to improved yield under heat stress regimes in potato [47]. Thermo tolerant alleles were introgressed into heat sensitive cultivar “Paymaster 404” from a donor accession “7456” of G. barbadense through backcross breeding [48]. A significant improvement in yield was realized under heat stress environment by adoption of gametic selection in maize [41]. A deep rooted cultivar “Nagina 22 (N22)” of aus rice exhibited high pollen viability and spikelet fertility (64–86%) under heat stress [49]. The thermo-tolerance of N22 was successfully introgressed into Xieqingzao B line through backcross method [50]. Dissecting out the genetic and physiological basis of thermo-tolerance will hasten up the development of resilient cultivars suited to hotter regions.
\nThe genetic basis of thermo-tolerance is not clearly understood because of complex trait inheritance. Advances in molecular approaches such as DNA marker identification and genotyping assay had paved way in determination of several QTL’s associated with high temperature tolerance [51]. In wheat, QTL’s were identified for canopy temperature, and chlorophyll fluorescence imparting tolerance to heat stress [52]. A major QTL “Htg 6.1” in lettuce was involved in enhancement of seed germination capacity at high temperature [53]. A recessive QTL for increased spikelet fertility under high temperature was dissected out in rice at chromosome 4. The identified QTL were found in several populations of heat tolerant rice cultivars [54]. Six QTL’s were involved to enhance fruit set at high temperature in tomato [55]. Five thermo tolerant QTL’s were identified in Brassica campestris by employing random amplified polymorphic DNA (RAPD) and amplified fragment length polymorphism (AFLP) markers [56]. In maize, eleven major QTL’s for increased pollen germination and pollen tube growth under high temperature was mapped using restriction fragment length polymorphism (RFLP) markers [57]. Identification of candidate QTL’s would pave way in precise introgression of heat tolerant genes into superior cultivars through marker assisted breeding approach.
\nThe closely associated markers with targeted QTL will hasten the recovery of superior genotypes with heat tolerant traits in a population. A marker assisted breeding approach was employed in rice to derive heat tolerant line with superior grain quality. Two flanking markers viz., ktIndel001 and RFT1 enclosing 1.5 Mb chromosomal region was transferred from tolerant cultivar “Kokoromachi” to Tohoku 168. Significant improvement in grain quality under high temperature was observed in the derived NIL’s compared to susceptible cultivar “Tohoku 168” [58]. Fourteen SSR markers linked to heat susceptibility index of grain filling per cent and single kernel weight was identified in bread wheat which was employed in marker assisted selection (MAS) to screen genotypes for thermo tolerance [59]. Utilization of MAS approach for heat tolerance remains less efficient because of high gene x environment and epistatic interactions. The low breeding efficiency can be resolved by genomic selection (GS) approach which involves wide number of molecular markers exhibiting high genome coverage. High genetic gain is realized in GS approach due to close association between predicted and true breeding value over generations [60].
\nAt present, transgenic approach also proves to be desirable tool for designing thermo tolerant lines via introgression of genes from diverse gene pools [61]. The genetic transformation was focused primarily on transcription factors, induction of heat shock proteins, molecular chaperones, osmolytes, antioxidant components and growth regulators [62]. Heat shock proteins play a primary role in imparting thermo tolerance in crop species. It is functionally associated with diverse group of molecular chaperones that is involved in restoration of degraded proteins to their native structure under high temperature. Induction of heat shock proteins through genetic manipulation was achieved in arabidopsis [63], maize [64], rice [65], soybean [66], and pepper [67]. The DREB gene family was also reported to impart heat tolerant response in many crop species. Over expression of ZmDREB2A in maize [68] and GmDREB2A in soybean [69] was associated with increased survival and adaptation under high temperature. Transgenic techniques were employed to alter membrane lipid properties for thermo-tolerance in crop species. High proportion of saturated fatty acid in membrane had increased tolerance under heat stress. Suppression of omega-3 fatty acid desaturase gene in chloroplast had reduced the accumulation of trieonic fatty acid in transgenic tobacco [70] and tomato [71] leading to thermo-tolerance. A significant accumulation of glycine betaine (osmolyte) was achieved in arabidopsis through transfer of “cod gene” from Arthrobacter globiformis [72]. High proportion of glycine betaine protects the PSII component by inhibiting the ROS activities under heat stress. Implementation of transgenic approaches in other crop species will accelerate the development of resilient genotypes suited to high temperature regimes.
\nDevelopment of thermo-tolerant lines has to be prioritized to meet out the future climatic change coupled with food demands. Knowledge on plant response and adaptation mechanisms to heat stress is required for framing out breeding strategies. It remains a challenging task in evolving resilient genotypes suited to high temperature because of less efficient screening protocols at field conditions. The existence of low genetic variation for heat response related traits limited the progress of conventional breeding approach in many crop species. Use of molecular breeding strategies had opened up several heat tolerant related QTL’s in crop species. However, still precise research work involving huge marker data is needed for attaining high breeding efficiency for thermo tolerance. Recently, the involvement of transgenic approach paved way for utilization of tolerant source from diverse gene pools. Study on induction of heat shock proteins led to increased thermo tolerance in many crop species. Similarly, other heat response related traits such as induction of antioxidant components, osmolytes, and chaperones were also included in transgenic approach for inducing heat stress tolerance. Thus, high economic yield could be realized at elevated temperature regimes with the involvement of combined breeding approaches.
\nThe authors are highly thankful to Dr. V. Geethalakshmi, Director, Directorate of Crop Management, Tamil Nadu Agricultural University (TNAU) for her valuable suggestions towards this chapter. We also acknowledge Dr. P. Jayamani, Professor and Head, Department of Pulses, TNAU; Dr. M. Raveendran, Professor and Head, Department of Biotechnology, TNAU; and Dr. K. Ganesamurthy, Professor and Head, Department of Rice, TNAU for rendering supportive documents on high temperature tolerance.
\nThe authors declare no conflict of interest towards this chapter.
The authors express their gratitude to the Directorate of Crop Management for providing scientific support on high temperature tolerance.
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