\\n\\n
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
\n'}],latestNews:[{slug:"intechopen-maintains-position-as-the-world-s-largest-oa-book-publisher-20201218",title:"IntechOpen Maintains Position as the World’s Largest OA Book Publisher"},{slug:"all-intechopen-books-available-on-perlego-20201215",title:"All IntechOpen Books Available on Perlego"},{slug:"oiv-awards-recognizes-intechopen-s-editors-20201127",title:"OIV Awards Recognizes IntechOpen's Editors"},{slug:"intechopen-joins-crossref-s-initiative-for-open-abstracts-i4oa-to-boost-the-discovery-of-research-20201005",title:"IntechOpen joins Crossref's Initiative for Open Abstracts (I4OA) to Boost the Discovery of Research"},{slug:"intechopen-hits-milestone-5-000-open-access-books-published-20200908",title:"IntechOpen hits milestone: 5,000 Open Access books published!"},{slug:"intechopen-books-hosted-on-the-mathworks-book-program-20200819",title:"IntechOpen Books Hosted on the MathWorks Book Program"},{slug:"intechopen-s-chapter-awarded-the-guenther-von-pannewitz-preis-2020-20200715",title:"IntechOpen's Chapter Awarded the Günther-von-Pannewitz-Preis 2020"},{slug:"suf-and-intechopen-announce-collaboration-20200331",title:"SUF and IntechOpen Announce Collaboration"}]},book:{item:{type:"book",id:"3424",leadTitle:null,fullTitle:"Food Industry",title:"Food Industry",subtitle:null,reviewType:"peer-reviewed",abstract:"Due to the increase in world population (more than seven billion inhabitants) the global food industry has the largest number of demanding and knowledgeable consumers. This population requires food products that fulfill the high quality standards established by the food industry organizations. Food shortages threaten human health, and also the disastrous extreme climatic events make food shortages even worse. This collection of articles is a timely contribution to issues relating to the food industry. The objective of this book is to provide knowledge appropriate for students, university researchers, and in general, for anyone wishing to obtain knowledge of food processing and to improve the food product quality.",isbn:null,printIsbn:"978-953-51-0911-2",pdfIsbn:"978-953-51-5345-0",doi:"10.5772/55834",price:159,priceEur:175,priceUsd:205,slug:"food-industry",numberOfPages:760,isOpenForSubmission:!1,isInWos:1,hash:"26d230385a4b7a517b44d60bf75e83de",bookSignature:"Innocenzo Muzzalupo",publishedDate:"January 16th 2013",coverURL:"https://cdn.intechopen.com/books/images_new/3424.jpg",numberOfDownloads:118142,numberOfWosCitations:132,numberOfCrossrefCitations:106,numberOfDimensionsCitations:304,hasAltmetrics:1,numberOfTotalCitations:542,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"May 9th 2012",dateEndSecondStepPublish:"May 30th 2012",dateEndThirdStepPublish:"August 26th 2012",dateEndFourthStepPublish:"September 25th 2012",dateEndFifthStepPublish:"December 4th 2012",currentStepOfPublishingProcess:5,indexedIn:"1,2,3,4,5,6",editedByType:"Edited by",kuFlag:!1,editors:[{id:"93139",title:"Dr.",name:"Innocenzo",middleName:null,surname:"Muzzalupo",slug:"innocenzo-muzzalupo",fullName:"Innocenzo Muzzalupo",profilePictureURL:"https://mts.intechopen.com/storage/users/93139/images/system/93139.png",biography:"Doctor Innocenzo Muzzalupo has received degree in “Biology” from the University of Calabria in 1993 and received his Ph.D. degree (1997) in “Chemistry” from the University of “La Sapienza” Rome. Currently, he is working as a member of Council for Agricultural Research and Economics, Research Centre for Olive, Citrus and Tree Fruit in Italy. After receiving his Ph.D. degree, he was appointed as post-doctoral researcher (1999) in “Food Science” at the University of Calabria. Between 1999 and 2008 he had a contract as professor of “Botany” at the University of Calabria. Following eight years of extensive research on olive characterization and on olive oil quality. His research areas include olive germplasm characterization, olive genes characterization, and analytical methods for olive oil traceability, olive oil quality. 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Neuroblastoma (NB) is one of the most common extracranial solid tumors of early childhood [1, 2]. Prognosis of patients with NB depends on tumor stage, patient’s age, and biologic feature of the tumor cells [3]. In patients under 1 year of age, NB is curable and sometimes spontaneously regress [4]. However, in older children with advanced stage, often the tumor is very aggressive, and patients have poor prognosis despite treatment with high-dose chemotherapy combined with autologous hematopoietic stem cell transplantation. Although immunotherapeutic therapy such as anti-GD2 monoclonal antibody has improved outcomes of advanced stage of NB, a number of patients still relapse and eventually die of disease [5]. The high incidence of resistance of high-risk NB to conventional therapies has prompted us to search for novel therapeutic approaches.
It was reported that phosphorylated protein kinase-B (Akt) correlates with poor patients’ prognosis in NB [6], and the phosphatidylinositol-3-kinase (PI3K)/Akt/mammalian target of rapamycin (mTOR) pathway has subsequently been linked to augmented cell survival [7] and increased resistance to chemotherapy in NB [8]. Therefore, targeting the PI3K/Akt/mTOR signaling pathway by appropriate inhibitors appears to be a promising strategy for overcoming therapy resistance [9].
Here, we demonstrate that NB cell lines are heterogeneous in their insulin growth factor-1 (IGF-1) receptor-mediated signaling [10]. The pattern of IGF-1 receptor/PI3K/Akt/mTOR pathway-mediated proliferation is an important determinant of the response to IGF-1 receptor antagonistic therapy in human NB [10]. Furthermore, our results highlight the potential of IGF-1 receptor/PI3K/Akt/mTOR signaling pathway as a promising target for NB treatment [10, 11, 12, 13].
IGF-1, IGF-2, and insulin belong to a family of mitogenic growth factors and are involved in normal growth and differentiation of most tissues. The biological actions of both IGFs and insulin can be mediated by the IGF-1 receptor which is involved in mitogenic, anti-apoptotic, and oncogenic transforming responses [14, 15]. The IGF-1 receptor has two extracellular α-subunits and two intracellular β-subunits that form a heterotetrameric complex. Ligand interaction with α-subunits triggers the autophosphorylation of tyrosine kinase domains within the β-subunit [16, 17, 18]. The tyrosine kinase domains are connected to several intracellular pathways such as PI3K/Akt [19, 20]. Dysregulation of the IGF-1 receptor pathway is involved in promoting oncogenic transformation, cell proliferation, metastasis, angiogenesis, and resistance in numerous malignant diseases, such as multiple myeloma [21], carcinomas [22], and NB [23]. Since high cellular heterogeneity is a hallmark of NB, which may account for the wide range of clinical presentations and nonuniform response to treatment, we hypothesized that NB cells are heterogeneous in their IGF-1 receptor signaling-mediated cell proliferation. Thirty-one NB cell lines were cultured in three different conditions, insulin-containing serum-free medium (SFM), RPMI1640 without FBS (serum starvation medium), and RPMI1640 with 10% FBS (serum-containing medium). Based on the response patterns, 31 cell lines were subdivided into three groups [10]. Group 1, which consisted of three NB cell lines, could proliferate for more than 3 days in SFM, RPMI1640 without FBS, and RPMI1640 with 10% FBS [10]. Group 2, which consisted of 10 cell lines, could proliferate in SFM and RPMI1640 with 10% FBS but not in RPMI1640 [10]. Group 3, which consisted of 18 NB cell lines, proliferated only in RPMI1640 with 10% FBS [10]. NB cell proliferation in RPMI1640 in the presence of exogenous IGF (IGF-1, IGF-2) and insulin was examined. These IGF and insulin accelerated cell proliferation in group 1 and group 2 NB cell lines but not in group 3 NB cell lines [10]. Group 1 NB cell lines were able to proliferate in RPMI1640 in the absence of exogenous IGF and insulin [10].
The IGF-1 receptor inhibitors, such as IGF-1 receptor-neutralizing antibodies and IGF-1 receptor antisense/siRNA, have been shown to block cancer cell proliferation [24]. Selective IGF-1 receptor inhibitor, picropodophyllin (PPP), lacks inhibitory activity on tyrosine phosphorylation of insulin receptor tyrosine kinase (RTK) [25]. Inhibition of the IGF-1 RTK with PPP is noncompetitive with respect to ATP, suggesting interference with the IGF-1 receptor at substrate level [26]. It is reported that PPP specifically blocks phosphorylation of the Tyr1136 residue in the activation loop of IGF-1 receptor kinase [27]. Inhibition of IGF-1 receptor with PPP has been demonstrated in a lot of cancers such as multiple myeloma [26], melanoma [28], breast cancer [29], and glioblastoma cells [30]. PPP inhibited Akt activation and suppressed cell proliferation in group 1 and group 2 NB cell lines but less in group 3 NB cell lines [10]. Elevation of ERK phosphorylation was only observed in group 1 NB cell lines [10]. U0126 is a MEK inhibitor effectively suppressed ERK phosphorylation, U0126 (2.5 μM) did not suppress cell proliferation induced by IGF-1, IGF-2, or insulin in group 1 and group 2 NB cell lines [10]. In groups 1 and 2 NB cell lines, IGF-1 receptor/PI3K/Akt pathway is critical for cell proliferation. Although IGF-1, IGF-2, and insulin activated Akt in group 3 NB cell line, cell proliferation was not increased in RPMI1640. This suggests that the activation of IGF-1R/Akt pathway is insufficient for cell proliferation [10]. Since IGF-1 and IGF-2 regulate apoptosis [31] and cell cycle progression [32], activation of caspase 3 and PARP was examined in NB cell lines. In group 1 NB cell lines, caspase 3 and PARP were not cleaved in RPMI1640 [10]. In group 2 NB cell lines, caspase 3 and PARP were cleaved in RPMI1640, and the cleavage of caspase 3 was suppressed by addition of IGF-1, IGF-2, and insulin [10]. In group 3 NB cell lines, cleavages of caspase 3 and PARP were observed in RPMI1640 [10]. However, they were not suppressed by IGF-1, IGF-2, and insulin, even though Akt activation was upregulated by IGF-1, IGF-2, and insulin [10]. Since PPP induced G2/M arrest and apoptosis by inhibiting IGF-1 receptor [26, 33], cell cycle phase distribution in NB cells was examined in the presence of PPP. PPP treatment (2.5 μM) for 12 hours increased the G2/M fraction and shifted cell cycle profile from G0-G1 dominant to G2/M dominant in group 1 and group 2 NB cell lines [10]. However, group 3 NB cell lines did not show G2/M arrest and the G0/G1 fraction was not affected [10]. Furthermore, cyclin B1, a marker protein of G2/M phase of cell cycle, was upregulated, and cyclin D1, a marker protein of G0/G1, synchronously declined in group 1 and group 2 NB cell lines [10]. However, accumulation of cyclin B1 and decline of cyclin D1 were not observed in group 3 NB cell lines [10]. This may be explained by the insensitiveness of group 3 cell lines to PPP-induced G2/M arrest.
MK2206 selectively inhibits AKT and has potency against Akt1 and Akt2 isoforms than Akt3. In pediatric solid tumors, MK2206 is effective in vitro and in vivo [34, 35]. In clinical trials, stable disease was observed in different kinds of cancers [36, 37]. Our results also suggested that MK2206 (2.5 μM) completely inhibited Akt phosphorylation and cell proliferation in NB cell lines [10]. Furthermore, MK2206 also impaired the cell proliferation induced by exogenous IGF and insulin in NB cell lines [10].
The PI3K/Akt/mTOR signaling cascade is one of the most important intracellular pathways, which is frequently activated in diverse cancers [38, 39]. The PI3K/Akt/mTOR pathway can be activated by transmembrane tyrosine kinase growth factor receptors, such as IGF-1 receptor, fibroblast growth factor receptors, ErbB family receptors, and others [40, 41]. Both mTOR S2448 and mTOR S2481 were extensively phosphorylated in NB cell lines [13]. Cell proliferation of NB cell lines was inhibited by AZD8055, a potent dual mTORC1-mTORC2 inhibitor [13]. According to the IC50, the NB cell lines were divided into two groups, sensitive to AZD8055 group (IC50 < 0.5 μM) and insensitive to AZD8055 group (IC50 > 0.5 μM) [13]. We also found that insensitive group showed lower mTOR (p < 0.001) expression and lower activity of mTOR complex 1 (p = 0.013) and mTOR complex 2 (p = 0.023) [13]. Cell cycle distribution analysis of NB cell lines was performed by flow cytometry. Cell cycle was affected with an increase in G0/G1 phase in dose-dependent manner of AZD8055 [13]. Western blotting analysis revealed that Cyclin D1 and Cyclin D3 were downregulated in AZD8055-treated NB cells [13]. AZD8055 inhibited both mTOR S2448 and mTOR S2481 phosphorylation significantly in a concentration-dependent manner [13]. Phosphorylation of downstream targets of mTOR complex 1, P70S6K T389 and 4E-BP1 S65, was also inhibited by AZD8055 treatment [13]. AZD8055 inhibited mTOR complex 2 substrates, Akt S473 and Akt T308 [13]. Although phosphorylation of Akt S473 was persistently inhibited in response to AZD8055 treatment, phosphorylation of Akt at the T308 site was inhibited for only 3–6 hours. These results indicate that AZD8055 inhibits mTOR activity and its downstream proteins in vitro in NB cells. Interestingly, NB cell lines were induced autophagy by AZD8055 treatment via downregulation of Akt/mTOR signaling pathway [13]. Autophagy inhibitor, 3-methyladenine, treatment resulted in a significant decrease of the AZD8055-induced apoptosis [13]. These results suggest that AZD8055 inhibited cell growth and induced cell cycle arrest, autophagy, and apoptosis in NB cells. Moreover, NB tumor growth in athymic nude mice was significantly inhibited by AZD8055 without toxicity [13]. Taken together, our results highlight that mTOR is a promising target for NB treatment. AZD8055 might be investigated for treatment of patients with advanced and refractory NB.
The problem that will plague single-target drugs is the cancer’s ability to activate alternative survival pathways leading to drug resistance and toxicity even in the multimodality setting. In addition, activation of multiple signaling pathways effectively causes cancer cell proliferation and survival. For example, RAS and PI3KCA are concurrently activated in melanoma, lung, and colorectal cancers [42, 43, 44]. These results suggested that combination therapies targeted on multiple signaling pathways could be more effective than targeting either pathway alone. Although AZD8055 is a promising drug in NB treatment [13], AZD8055-resistant NB sublines were acquired by prolonged stepwise exposure. After incubation with AZD8055 for 4–12 weeks, two acquired AZD8055-resistant sublines proliferated stably in RPMI1640 plus 10% FBS medium in the presence of AZD8055 (3 μM) [12]. The AZD8055-resistant sublines exhibited marked resistance to AZD8055 compared to the parent cells, and IC50 of the resistant sublines was 60–100 times higher than the parent NB cell lines [12]. By cell cycle analysis, accumulation of S phase was observed, and cyclin D3 and CDK4 were upregulated in AZD8055-resistant sublines [12]. Although AZD8055 treatment inhibited MEK/ERK activation in parent cells, MEK/ERK phosphorylation was continued despite AZD8055 treatment in resistant cells [12]. The combination therapy of AZD8055 and MEK/ERK inhibitor U0126 significantly inhibited cell proliferation compared to U0126 monotherapy [12], suggesting that combination therapy can overcome AZD8055 resistance. Furthermore, in athymic mice model, AZD8055 and U0126 co-treatment was more efficient to suppress resistant NB tumor growth compared to U0126 monotherapy [12].
MK2206 treatment induced a dose-dependent inhibition of cell proliferation, with IC50 ranging from 1.22 to 4.35 μM in NB cell lines [11]. MK2206-resistant cells were induced by stepwise escalation of MK-2206 exposure (4–12 weeks) [11]. These cells proliferated in RPMI1640 plus 10% FBS medium in the presence of MK2206 (5 μM), while cell death was induced in parent cells [11]. IC50 of the resistant cell lines was 6–7 times higher than the parent NB cell lines [11]. Small-molecule GSK2334470 selectively inhibits 3-phosphoinositide-dependent protein kinase 1 (PDK1) with low concentration but does not suppress the activity of other protein kinases at higher concentrations [45]. Although GSK2334470 attenuated cell proliferation in both parent cells and MK2206-resistant sublines, IC50 of GSK2334470 in resistant sublines was lower than that of parent cell lines [11]. GSK2334470 induced G0-G1 accumulation of cell cycle phase distribution in parent cell lines [11]. In MK2206-resistant sublines, G0-G1 accumulation induced by GSK2334470 was higher than parent cell lines [11].
Amplification of the MYCN oncogene is the most powerful single predictor of adverse outcome of NB [46]. MYCN amplification is observed in about 20% of all NB patients and is usually associated with fatal outcome of the disease [47]. Schramm et al. demonstrated transcriptomal upregulation of mTOR-related genes by MYCN [48]. MYCN-driven NB in mice displayed activation of the mTOR pathway on the protein level, and activation of MYCN in NB cells resulted in high sensitivity toward mTOR inhibition [48]. Therefore, it is examined whether MYCN status of NB cell lines affects susceptibility to AZD8055. Using fluorescence in situ hybridization (FISH), we observed MYCN amplification in all investigated nuclei of NB19, IMR32, TGW, OZAWA, LAN-1, SMS-KAN, and SCMC-N4 cell lines (Figure 1A). As shown in Figure 1A, MYCN amplification was not observed in INDEN (loss/imbalance), SK-N-SH (gain), KP-N-SI (loss/imbalance), KP-N-SIFA (gain), and SJ-N-KP (loss/imbalance). As shown in Figure 1B, the relationship between MYCN status and susceptibility to AZD8055 was not found.
Effect of MYCN status on susceptibility to AZD8055. (A) MYCN gene in tested 12 NB cell lines. MYCN status was defined as previously [53]. No alteration: cells with two MYCN signals and two CEP2 signals; amplification: the number of MYCN signals is at least 10 copies greater than the control probe signals; loss/imbalance: presence of at least two MYCN signals and increased CEP2 signals; gain: the number of MYCN signals is 1–9 copies more than CEP2 signals. (B) Fifteen NB cell lines were treated with AZD8055 at different concentrations in RPMI1640 + 10% FBS. Cell growth was evaluated as cell numbers at 72 hours. Data are expressed as the mean ± SD. IC50 (half maximal inhibitory concentration) of the 15 NB cell lines was calculated depending on the MTT results.
Constitutive activation of the ALK receptor tyrosine kinase by mutation or translocation appears to contribute to the malignant phenotype of several cancers, including NB, making it a potentially therapeutic target [49]. Both wild type ALK and the F1174L-mutated ALK upregulate MYCN expression [50], and these two genes lead to constitutive phosphorylation of ALK and of downstream signaling molecules, PI3K/AKT/mTOR, that are critical for cell proliferation and survival [51]. ALKF1174L cosegregates with MYCN amplification in patients, and this combination is associated with a particularly poor prognosis, as shown by the fatal outcome of 9 of 10 children with ALKF1174L/MYCN-amplified tumors [52]. Targeting ALK with PI3K/Akt/mTOR inhibitors may inhibit cell growth in tumor lines with concomitant MYCN amplification.
NB cell lines can be categorized into three groups by the patterns of IGF-1R/Akt pathway response. PPP, MK2206, and AZD8055 showed significant antitumor effect in NB cells. Our current results highlight the potential of IGF-1R/PI3K/Akt/mTOR pathway as a promising target for NB treatment. PPP, MK2206, and AZD8055 should be further investigated for NB treatment in clinical trials.
The authors declare no potential conflicts of interest.
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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