The yield losses in cereal crops because of abiotic stress and the expected huge losses from climate change indicate our urgent need for useful traits to achieve food security. The stay-green (SG) is a secondary trait that enables crop plants to maintain their green leaves and photosynthesis capacity for a longer time after anthesis, especially under drought and heat stress conditions. Thus, SG plants have longer grain-filling period and subsequently higher yield than non-SG. SG trait was recognized as a superior characteristic for commercially bred cereal selection to overcome the current yield stagnation in alliance with yield adaptability and stability. Breeding for functional SG has contributed in improving crop yields, particularly when it is combined with other useful traits. Thus, elucidating the molecular and physiological mechanisms associated with SG trait is maybe the key to defeating the stagnation in productivity associated with adaptation to environmental stress. This review discusses the recent advances in SG as a crucial trait for genetic improvement of the five major cereal crops, sorghum, wheat, rice, maize, and barley with particular emphasis on the physiological consequences of SG trait. Finally, we provided perspectives on future directions for SG research that addresses present and future global challenges.

Stay-green (sometimes staygreen) refers to the heritable delayed foliar senescence character in model and crop plant species. In a cosmetic stay-green, a lesion interferes with an early step in chlorophyll catabolism. The possible contribution of synthesis to chlorophyll turnover in cosmetic stay-greens is considered. In functional stay-greens, the transition from the carbon capture period to the nitrogen mobilization (senescence) phase of canopy development is delayed, and/or the senescence syndrome proceeds slowly. Yield and composition in high-carbon (C) crops such as cereals, and in high-nitrogen (N) species such as legumes, reflect the source-sink relationship with canopy C capture and N remobilization. Quantitative trait loci studies show that functional stay-green is a valuable trait for improving crop stress tolerance, and is associated with the domestication syndrome in cereals. Stay-green variants reveal how autumnal senescence and dormancy are coordinated in trees. The stay-green phenotype can be the result of alterations in hormone metabolism and signalling, particularly affecting networks involving cytokinins and ethylene. Members of the WRKY and NAC families, and an ever-expanding cast of additional senescence-associated transcription factors, are identifiable by mutations that result in stay-green. Empirical selection for functional stay-green has contributed to increasing crop yields, particularly where it is part of a strategy that also targets other traits such as sink capacity and environmental sensitivity and is associated with appropriate crop management methodology. The onset and progress of senescence are phenological metrics that show climate change sensitivity, indicating that understanding stay-green can contribute to the design of appropriate crop types for future environments. © The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.

Stay-green mutants are delayed in leaf senescence and have been identified from different plant species, including many crops. Functional stay-greens have the potential to increase plant productivity. In cosmetic stay-greens, however, retention of chlorophyll during senescence is uncoupled from a decline of photosynthetic capacity in these mutants. For many cosmetic stay-green mutants, including Gregor Mendel's famous green cotyledon pea variety, molecular defects were recently identified in orthologous stay-green genes. Stay-green genes encode members of a new family of chloroplast-located proteins, which are likely to function in dismantling of photosynthetic chlorophyll-apoprotein complexes. Their activity is considered as a prerequisite for both chlorophyll and apoprotein degradation during senescence.

Production of sorghum [Sorghum bicolor (L.) Moench], an important cereal crop in semiarid regions of the world, is often limited by drought. When water is limiting during the grain‐filling period, hybrids possessing the stay‐green trait maintain more photosynthetically active leaves than hybrids not possessing this trait. To improve yield under drought, knowledge of the extent of genetic variation in green leaf area retention is required. Field studies were undertaken in northeastern Australia on a cracking and self‐mulching gray clay to determine the effects of water regime and hybrid on the components of green leaf area at maturity (GLAM). Nine hybrids varying in stay‐green were grown under a fully irrigated control, postflowering water deficit, and terminal (pre‐ and postflowering) water deficit. Water deficit reduced GLAM by 67% in the terminal drought treatment compared with the fully irrigated control. Under terminal water deficit, hybrids possessing the B35 and KS19 sources of stay‐green retained more GLAM (1260 cm2 plant−1) compared with intermediate (780 cm2 plant−1) and senescent (670 cm2 plant−1) hybrids. RQL12 hybrids (KS19 source of stay‐green) displayed delayed onset and reduced rate of senescence; A35 hybrids displayed only delayed onset. Visual rating of green leaf retention was highly correlated with measured GLAM, although this procedure is constrained by an inability to distinguish among the functional mechanisms determining the phenotype. Linking functional rather than phenotypic differences to molecular markers may improve the efficiency of selecting for traits such as stay‐green.

Plants synchronize their life history events with proper seasonal conditions, and as the fitness consequences of each life stage depend on previous and/or subsequent one, changes in environmental cues create cascading effects throughout their whole life cycle. For monocarpic plants, proper senescence timing is very important as the final production of plants depends on it. Citing available literatures, this review discusses how plants not only may delay senescence until after they reproduce successfully, but they may also bring senescence time forward, in order to reproduce in favored conditions. It demonstrates that even though senescence is part of aging, it does not necessarily mean plants have to reach a certain age to senesce. Experiments using different aged plants have suggested that in interest of their final outcome and fitness, plants carefully weigh out environmental cues and transit to next developmental phase at proper time, even if that means transiting to terminal senescence phase earlier and shortening their lifespan. How much plants have control over senescence timing and how they balance internal and external signals for that is not well understood. Future studies are needed to identify processes that trigger senescence timing in response to environment and investigate genetic/epigenetic mechanisms behind it.

Environmental stresses from climate change can alter source-sink relations during plant maturation, leading to premature senescence and decreased yields. Elucidating the genetic control of natural variations in senescence in wheat (Triticum aestivum) can be accelerated using recent developments in unmanned aerial vehicle (UAV) based imaging techniques. Here, we describe the use of UAVs to quantify senescence in wheat using vegetative indices (VIs) derived from multispectral images. We detected senescence with high heritability, as well as its impact on grain yield, in a doubled-haploid population and parent cultivars at various growth time points after anthesis in the field. Selecting for slow senescence using a combination of different UAV-based VIs was more effective than using a single ground-based vegetation index. We identified 28 quantitative trait loci (QTLs) for vegetative growth, senescence, and grain yield using a 660K single-nucleotide polymorphism (SNP) array. Seventeen of these new QTLs for VIs from UAV-based multispectral imaging were mapped on chromosomes 2B, 3A, 3D, 5A, 5D, 5B, and 6D; these QTLs have not been reported previously using conventional phenotyping methods. This integrated approach allowed us to identify an important, previously unreported, senescence-related locus on chromosome 5D that showed high phenotypic variation (up to 18.1%) for all UAV-based VIs at all time points during grain filling. This QTL was validated for slow senescence by developing kompetitive allele-specific PCR (KASP) markers in a natural population. Our results suggest that UAV-based high-throughput phenotyping is advantageous for temporal assessment of the genetics underlying senescence in wheat.© The Author(s) 2021. Published by Oxford University Press on behalf of American Society of Plant Biologists.

Delayed senescence, or stay-green, contributes to a longer grain-filling period and has been regarded as a desirable characteristic for the production of a number of crops including wheat. In the present study, in order to identify quantitative trait loci (QTLs) for traits related to the progression of wheat flag leaf senescence, green leaf area duration (GLAD) of a doubled haploid (DH) population, derived from two winter wheat varieties Hanxuan10 and Lumai14, was visually estimated under two water conditions and was recorded at 3-day intervals from 10 days after anthesis to physiological maturity using a 0–9 scale. According to GLAD, parameters related to the progression of senescence of DH lines and their parents were estimated by the Gompertz statistical model. Based on the model parameters, DH lines were categorized into three groups under drought stress and four groups under well-watered conditions. A total of 24 additive QTLs and 23 pairs of epistatic QTLs for parameters related to the progression of senescence were identified on 18 chromosomes, except for 3B, 1D and 6D. Of the QTLs detected, 14 and 10 additive QTLs were associated with the investigated traits under drought stress and well-watered conditions, respectively. Furthermore, 4, 7, 6, 2 and 2 additive QTLs for traits related to progression of senescence were clustered around the same or similar regions of chromosomes 1A, 1B, 5A, 5B and 7A, respectively. The present data provided the genetic basis for high phenotypic correlations among traits related to the progression of wheat flag leaf senescence. In addition, 17 loci were co-located or linked with previously reported QTLs regulating chlorophyll fluorescence, high-light-induced photo-oxidation, or heat stress and dark-induced senescence. The markerXwmc336on chromosome 1A, responsible for the onset and end times of leaf senescence, the time to maximum rate of senescence, the time to reach 75% senescence and chlorophyll content under drought stress may be helpful for marker-assisted selection breeding of wheat.

A restricted range in height and phenology of the elite Seri/Babax recombinant inbred line (RIL) population makes it ideal for physiological and genetic studies. Previous research has shown differential expression for yield under water deficit associated with canopy temperature (CT). In the current study, 167 RILs plus parents were phenotyped under drought (DRT), hot irrigated (HOT), and temperate irrigated (IRR) environments to identify the genomic regions associated with stress-adaptive traits. In total, 104 QTL were identified across a combination of 115 traits × 3 environments × 2 years, of which 14, 16, and 10 QTL were associated exclusively with DRT, HOT, and IRR, respectively. Six genomic regions were related to a large number of traits, namely 1B-a, 2B-a, 3B-b, 4A-a, 4A-b, and 5A-a. A yield QTL located on 4A-a explained 27 and 17% of variation under drought and heat stress, respectively. At the same location, a QTL explained 28% of the variation in CT under heat, while 14% of CT variation under drought was explained by a QTL on 3B-b. The T1BL.1RS (rye) translocation donated by the Seri parent was associated with decreased yield in this population. There was no co-location of consistent yield and phenology or height-related QTL, highlighting the utility of using a population with a restricted range in anthesis to facilitate QTL studies. Common QTL for drought and heat stress traits were identified on 1B-a, 2B-a, 3B-b, 4A-a, 4B-b, and 7A-a confirming their generic value across stresses. Yield QTL were shown to be associated with components of other traits, supporting the prospects for dissecting crop performance into its physiological and genetic components in order to facilitate a more strategic approach to breeding.

Leaf aging is a highly regulated developmental process, which is also influenced profoundly by diverse environmental conditions. Accumulating evidence in recent years supports that plant responsiveness to abiotic stress is intimately related with leaf longevity. However, molecular mechanisms underlying the signaling crosstalks and regulatory schemes are yet unknown. In this work, we demonstrate that an abscisic acid (ABA)–responsive NAC transcription factor VND-INTERACTING2 (VNI2) integrates ABA-mediated abiotic stress signals into leaf aging by regulating a subset of COLD-REGULATED (COR) and RESPONSIVE TO DEHYDRATION (RD) genes. The VNI2 gene was induced by high salinity in an ABA-dependent manner. In addition, spatial and temporal expression patterns of the VNI2 gene are correlated with leaf aging and senescence. Accordingly, leaf aging was delayed in transgenic plants overexpressing the VNI2 gene but significantly accelerated in a VNI2-deficient mutant. The VNI2 transcription factor regulates the COR and RD genes by binding directly to their promoters. Notably, transgenic plants overexpressing the COR or RD genes exhibited prolonged leaf longevity. These observations indicate that the VNI2 transcription factor serves as a molecular link that integrates plant responses to environmental stresses into modulation of leaf longevity.

Leaf senescence is an essential physiological process in plants that supports the recycling of nitrogen and other nutrients to support the growth of developing organs, including young leaves, seeds, and fruits. Thus, the regulation of senescence is crucial for evolutionary success in wild populations and for increasing yield in crops. Here, we describe the influence of a NAC transcription factor, SlNAP2 ( NAC-like, activated by Apetala3/Pistillata), that controls both leaf senescence and fruit yield in tomato (). expression increases during age-dependent and dark-induced leaf senescence. We demonstrate that SlNAP2 activates (), a homolog of Arabidopsis (), chlorophyll degradation genes such as ( senescence-inducible chloroplast stay-green protein 1) and ( pheide oxygenase), and other downstream targets by directly binding to their promoters, thereby promoting leaf senescence. Furthermore, SlNAP2 directly controls the expression of genes important for abscisic acid (ABA) biosynthesis, 9-cis-epoxycarotenoid dioxygenase 1 (); transport, ABC transporter G family member 40 (); and degradation, ABA 8'-hydroxylase (), indicating that SlNAP2 has a complex role in establishing ABA homeostasis during leaf senescence. Inhibiting expression in transgenic tomato plants impedes leaf senescence but enhances fruit yield and sugar content likely due to prolonged leaf photosynthesis in aging tomato plants. Our data indicate that SlNAP2 has a central role in controlling leaf senescence and fruit yield in tomato.© 2018 American Society of Plant Biologists. All rights reserved.

Modern tomato (Solanum lycopersicum) varieties are bred for uniform ripening (u) light green fruit phenotypes to facilitate harvests of evenly ripened fruit. U encodes a Golden 2-like (GLK) transcription factor, SlGLK2, which determines chlorophyll accumulation and distribution in developing fruit. In tomato, two GLKs--SlGLK1 and SlGLK2--are expressed in leaves, but only SlGLK2 is expressed in fruit. Expressing GLKs increased the chlorophyll content of fruit, whereas SlGLK2 suppression recapitulated the u mutant phenotype. GLK overexpression enhanced fruit photosynthesis gene expression and chloroplast development, leading to elevated carbohydrates and carotenoids in ripe fruit. SlGLK2 influences photosynthesis in developing fruit, contributing to mature fruit characteristics and suggesting that selection of u inadvertently compromised ripe fruit quality in exchange for desirable production traits.

Fruit ripening is the summation of changes rendering fleshy fruit tissues attractive and palatable to seed dispersing organisms. For example, sugar content is influenced by plastid numbers and photosynthetic activity in unripe fruit and later by starch and sugar catabolism during ripening. Tomato fruit are sinks of photosynthate, yet unripe green fruit contribute significantly to the sugars that ultimately accumulate in the ripe fruit. Plastid numbers and chlorophyll content are influenced by numerous environmental and genetic factors and are positively correlated with photosynthesis and photosynthate accumulation. GOLDEN2-LIKE (GLK) transcription factors regulate plastid and chlorophyll levels. Tomato (Solanum lycopersicum), like most plants, contains two GLKs (i.e., GLK1 and GLK2/UNIFORM). Mutant and transgene analysis demonstrated that these genes encode functionally similar peptides, though differential expression renders GLK1 more important in leaves, while GLK2 is predominant in fruit. A latitudinal gradient of GLK2 expression influences the typical uneven coloration of green and ripe wild-type fruit. Transcriptome profiling revealed a broader fruit gene expression gradient throughout development. The gradient influenced general ripening activities beyond plastid development and was consistent with the easily observed yet poorly studied ripening gradient present in tomato and many fleshy fruits.

Stay-green plants retain green leaves longer after anthesis and can have improved yield, particularly under water limitation. As senescence is a dynamic process, genotypes with different senescence patterns may exhibit similar final normalised difference vegetative index (NDVI). By monitoring NDVI from as early as awn emergence to maturity, we demonstrate that analysing senescence dynamics improves insight into genotypic stay-green variation. A senescence evaluation tool was developed to fit a logistic function to NDVI data and used to analyse data from three environments for a wheat (Triticum aestivum L.) population whose lines contrast for stay-green. Key stay-green traits were estimated including, maximum NDVI, senescence rate and a trait integrating NDVI variation after anthesis, as well as the timing from anthesis to onset, midpoint and conclusion of senescence. The integrative trait and the timing to onset and mid-senescence exhibited high positive correlations with yield and a high heritability in the three studied environments. Senescence rate was correlated with yield in some environments, whereas maximum NDVI was associated with yield in a drought-stressed environment. Where resources preclude frequent measurements, we found that NDVI measurements may be restricted to the period of rapid senescence, but caution is required when dealing with lines of different phenology. In contrast, regular monitoring during the whole period after flowering allows the estimation of senescence dynamics traits that may be reliably compared across genotypes and environments. We anticipate that selection for stay-green traits will enhance genetic progress towards high-yielding, stay-green germplasm.

Germplasm, genetics, phenotyping, and selection, combined with a clear definition of product targets, are the foundation of successful hybrid maize breeding. Breeding maize hybrids with superior yield for the drought-prone regions of the US corn-belt involves integration of multiple drought-specific technologies together with all of the other technology components that comprise a successful maize hybrid breeding programme. Managed-environment technologies are used to enable scaling of precision phenotyping in appropriate drought environmental conditions to breeding programme level. Genomics and other molecular technologies are used to study trait genetic architecture. Genetic prediction methodology was used to breed for improved yield performance for drought-prone environments. This was enabled by combining precision phenotyping for drought performance with genetic understanding of the traits contributing to successful hybrids in the target drought-prone environments and the availability of molecular markers distributed across the maize genome. Advances in crop growth modelling methodology are being used to evaluate the integrated effects of multiple traits for their combined effects and evaluate drought hybrid product concepts and guide their development and evaluation. Results to date, lessons learned, and future opportunities for further improving the drought tolerance of maize for the US corn-belt are discussed. © The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.

Stay-green sorghum plants exhibit greener leaves and stems during the grain-filling period under water-limited conditions compared with their senescent counterparts, resulting in increased grain yield, grain mass, and lodging resistance. Stay-green has been mapped to a number of key chromosomal regions, including Stg1, Stg2, Stg3, and Stg4, but the functions of these individual quantitative trait loci (QTLs) remain unclear. The objective of this study was to show how positive effects of Stg QTLs on grain yield under drought can be explained as emergent consequences of their effects on temporal and spatial water-use patterns that result from changes in leaf-area dynamics. A set of four Stg near-isogenic lines (NILs) and their recurrent parent were grown in a range of field and semicontrolled experiments in southeast Queensland, Australia. These studies showed that the four Stg QTLs regulate canopy size by: (1) reducing tillering via increased size of lower leaves, (2) constraining the size of the upper leaves; and (3) in some cases, decreasing the number of leaves per culm. In addition, they variously affect leaf anatomy and root growth. The multiple pathways by which Stg QTLs modulate canopy development can result in considerable developmental plasticity. The reduction in canopy size associated with Stg QTLs reduced pre-flowering water demand, thereby increasing water availability during grain filling and, ultimately, grain yield. The generic physiological mechanisms underlying the stay-green trait suggest that similar Stg QTLs could enhance post-anthesis drought adaptation in other major cereals such as maize, wheat, and rice. © The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology.

Stay-green is an integrated drought adaptation trait characterized by a distinct green leaf phenotype during grain filling under terminal drought. We used sorghum (Sorghum bicolor), a repository of drought adaptation mechanisms, to elucidate the physiological and genetic mechanisms underpinning stay-green. Near-isogenic sorghum lines (cv RTx7000) were characterized in a series of field and managed-environment trials (seven experiments and 14 environments) to determine the influence of four individual stay-green (Stg1-4) quantitative trait loci (QTLs) on canopy development, water use and grain yield under post-anthesis drought. The Stg QTL decreased tillering and the size of upper leaves, which reduced canopy size at anthesis. This reduction in transpirational leaf area conserved soil water before anthesis for use during grain filling. Increased water uptake during grain filling of Stg near-isogenic lines (NILs) relative to RTx7000 resulted in higher post-anthesis biomass production, grain number and yield. Importantly, there was no consistent yield penalty associated with the Stg QTL in the irrigated control. These results establish a link between the role of the Stg QTL in modifying canopy development and the subsequent impact on crop water use patterns and grain yield under terminal drought. © 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.

Nitrogen (N), as a macro-element, plays a vital role in plant growth and development. N deficiency affects plant productivity by decreasing photosynthesis, leaf area and longevity of green leaf. To date, many studies have reported that the relationship between photosynthesis and N supply. Here, we summarized the physiological response of photosynthesis to N deficiency in leaf structure and N allocation within the leaf. In serious N stress, photosynthetic rate decreases for almost all plants. The reasons as follows:(1) reducing stomatal conductance of mesophyll cell (g) and bundle sheath cells (g) which influences intercellular CO concentration; (2) reducing the content of bioenergetics and light-harvesting protein which inhibits electron transport rate and increase the light energy dissipated as heat; (3) reducing the content and/or activity of photosynthetic enzymes which reduces carboxylation rate. During reproductive stage, N stress induces plant senescence and N components degradation, especially photosynthetic enzymes and thylakoid N, and thus reduces photosynthesis. To keep high grain yield in low N deficiency, we should choose the genotype with higher N allocation within bioenergetics and lower degradation of photosynthetic enzymes. This review provides a generalized N allocation in response to N stress and gives a new prospect for breeding N-efficient genotypes.Copyright © 2020 Elsevier Masson SAS. All rights reserved.

Sorghum [Sorghum bicolor (L.) Moench] hybrids containing the stay‐green trait retain more photosynthetically active leaves under drought than do hybrids that do not contain this trait. Since the longevity and photosynthetic capacity of a leaf are related to its N status, it is important to clarify the role of N in extending leaf greenness in stay‐green hybrids. Field studies were conducted in northeastern Australia to examine the effect of three water regimes and nine hybrids on N uptake and partitioning among organs. Nine hybrids varying in the B35 and KS19 sources of stay‐green were grown under a fully irrigated control, post‐flowering water deficit, and terminal water deficit. For hybrids grown under terminal water deficit, stay‐green was viewed as a consequence of the balance between N demand by the grain and N supply during grain filling. On the demand side, grain numbers were 16% higher in the four stay‐green than in the five senescent hybrids. On the supply side, age‐related senescence provided an average of 34 and 42 kg N ha−1 for stay‐green and senescent hybrids, respectively. In addition, N uptake during grain filling averaged 116 and 82 kg ha−1 in stay‐green and senescent hybrids. Matching the N supply from these two sources with grain N demand found that the shortfall in N supply for grain filling in the stay‐green and senescent hybrids averaged 32 and 41 kg N ha−1, resulting in more accelerated leaf senescence in the senescent hybrids. Genotypic differences in delayed onset and reduced rate of leaf senescence were explained by differences in specific leaf nitrogen and N uptake during grain filling. Leaf nitrogen concentration at anthesis was correlated with onset and rate of leaf senescence under terminal water deficit.

The expression of stay-green (SG) characteristic in sorghum under water stress was related to N supply. SG genotype performed better than a non-stay-green (NSG) genotype at medium and high N levels. The differences in physiological parameters between SG and NSG genotypes were not significant at low N level and severe water stress. Grain sorghum [Sorghum bicolor (L.) Moench] with stay-green (SG) trait has the potential to produce more biomass and use soil water and nitrogen (N) more efficiently under post-flowering water stress. Previous studies were mostly conducted without N deficiency and more information is needed for interactions among soil N availability, SG genotype, and post-flowering water stress. In this study, the differences in leaf growth and senescence, shoot and root biomass, evapotranspiration (ET), water use efficiency (WUE), leaf photosynthetic responses, and nitrogen use efficiency (NUE) between a SG genotype (BTx642) and a non-stay-green (NSG) genotype (Tx7000) were examined. The two genotypes were grown at three N levels (Low, LN; Medium, MN; High, HN) and under three post-flowering water regimes (No water deficit, ND; Moderate water deficit, MD; Severe water deficit, SD). The genotypic difference was generally significant while it frequently interacted with N levels and water regimes. At medium and high N levels, SG genotype consistently had greater green leaf area, slower senescence rate, more shoot biomass and root biomass, and greater WUE and NUE than the NSG genotype under post-flowering drought. However, differences in several variables (e.g., leaf senescence, ET, WUE and NUE) between genotypes were not significant under SD at LN. At HN and MN, photosynthetic function of SG genotype was better maintained under drought. At LN, SG genotype maintained greater green leaf area but had lower photosynthetic activity than the NSG genotype. Nonetheless, adequate N supply is important for SG genotype under drought and greater root biomass may contribute to greater NUE in SG genotype.© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.

试验选用持绿型冬小麦(Triticum aestivum) ‘豫麦66’ (‘Ym66’)和‘潍麦8号’ (‘Wm8’)为研究材料, 以当地生产上起主导作用的冬小麦品种‘小偃22’ (‘XY22’)和‘小偃6号’ (‘XY6’)为对照。花后用塑料薄膜搭建成增温棚进行高温处理, 测定各品种绿叶数目、叶绿素和丙二醛(MDA)含量及叶片细胞膜透性, 并研究籽粒灌浆成熟期高温对持绿型小麦籽粒淀粉合成相关酶及粒重的影响。结果表明, 高温处理后, 各品种的绿叶数目和叶绿素含量都减少, MDA含量和膜透性都增加, 说明高温加速了小麦叶片衰老。同时, 各品种籽粒中与淀粉合成相关的酶(蔗糖合成酶(SS)和腺苷二磷酸葡萄糖焦磷酸化酶(AGPP)、可溶性淀粉合酶(SSS))活性都低于正常生长下的籽粒中的酶活性, 其中高温对籽粒SS和AGPP活性的影响不显著,而对籽粒SSS活性的影响显著(p = 0.015)。品种间比较, 持绿型小麦在两种处理下, 都表现出较多的绿叶数目和较高的叶绿素含量; 且3种与淀粉合成相关的酶活性也都高于非持绿型小麦, 说明持绿型小麦酶活性受高温抑制程度较小。相关性分析表明, 所有品种籽粒SS、AGPP、SSS活性都与籽粒灌浆速率成极显著的正相关(相关系数r分别为0.905、0.419和0.801)。因而, 持绿型小麦不仅具有较好的持绿特性, 而且籽粒中与淀粉合成相关的3种酶活性都较高, 这有利于其籽粒淀粉的合成, 从而增加籽粒产量。

The stay-green leaf phenotype is typically associated with increased yields and improved stress resistance in maize breeding, due to higher nitrogen (N) nutrient levels that prolong greenness. The application of N fertilizer can regulate the N status of plants, and furthermore, impact the photosynthetic rates of leaves at the productive stage; however, N deficiencies and N excesses will reduce maize yields. Consequently, it is necessary to develop N fertilizer management strategies for different types of stay-green maize. For this study, the senescent cultivar Lianchuang 808 (LC808), moderate-stay-green cultivar Zhengdan 958 (ZD958), and over stay-green cultivar Denghai 685 (DH685) were selected as experimental models. Our results revealed that yields of ZD958 were slightly higher than DH685 and notably improved over than LC808. Compared with a non-stay-green cultivar LC808, ZD958 and DH685 still maintained higher chlorophyll contents and cell activities following the silking stage, while efficiently slowing the senescence rate. The supply of N fertilizer significantly prolonged leaf greenness and delayed senescence for ZD958 and DH685; however, the effect was not obvious for LC808. The stem remobilization efficiency of N was higher in the moderate-stay-green cultivar ZD958, in contrast to LC808, while the transfer of leaf N was lower than LC808, which guaranteed high leaf N levels, and that sufficient N was transferred to grains in ZD958. To obtain the highest yields, the optimal N fertilizer rates were 228.1 kg hm−2 for LC0808, 180 kg hm−2 for ZD958, and 203.8 kg hm−2 for DH685. In future, the selection of stay-green type crops might serve as an important agricultural strategy to reduce the quantity of N fertilizer and increase N efficiency.

Senescence is a developmental process which in annual crop plants overlaps with the reproductive phase. Senescence might reduce crop yield when it is induced prematurely under adverse environmental conditions. This review covers the role of senescence for the productivity of crop plants. With the aim to enhance productivity, a number of functional stay-green cultivars have been selected by conventional breeding, in particular of sorghum and maize. In many cases, a positive correlation between leaf area duration and yield has been observed, although in a number of other cases, stay-green cultivars do not display significant effects with regards to productivity. In several crops, the stay-green phenotype is observed to be associated with a higher drought resistance and a better performance under low nitrogen conditions. Among the approaches used to achieve stay-green phenotypes in transgenic plants, the expression of the IPT gene under control of senescence-associated promoters has been the most successful. The promoters employed for senescence-regulated expression contain cis-elements for binding of WRKY transcription factors and factors controlled by abscisic acid. In most crops transformed with such constructs the stay-green character has led to increased biomass, but only in few cases to increased seed yield. A coincidence of drought stress resistance and stay-green trait is observed in many transgenic plants.

Soil acidification is a major problem in soils of intensive Chinese agricultural systems. We used two nationwide surveys, paired comparisons in numerous individual sites, and several long-term monitoring-field data sets to evaluate changes in soil acidity. Soil pH declined significantly (P < 0.001) from the 1980s to the 2000s in the major Chinese crop-production areas. Processes related to nitrogen cycling released 20 to 221 kilomoles of hydrogen ion (H+) per hectare per year, and base cations uptake contributed a further 15 to 20 kilomoles of H+ per hectare per year to soil acidification in four widespread cropping systems. In comparison, acid deposition (0.4 to 2.0 kilomoles of H+ per hectare per year) made a small contribution to the acidification of agricultural soils across China.

The association between postflowering drought tolerance and the stay‐green phenotype in sorghum [Sorghum bicolor (L.) Moench] is well established. Recent studies link the stay‐green phenotype with levels of dhurrin and soluble sugars in the leaves. Postflowering drought response has always been associated with stalk rot resistance, especially charcoal rot. However, apart from the common knowledge that charcoal rot requires a dry environment, the mechanism governing these associations is not clear. The objectives of this study were to (i) determine the effects of major stay‐green (stg) quantitative trait loci (QTL) on response to infection by two stalk rot pathogens, Macrophomina phaseolina and Fusarium thapsinum, and (ii) examine the possible relationship between levels of dhurrin and soluble sugar in the leaves and response to stalk rot diseases. Fourteen genotypic groups derived from the Tx642 × Tx7000 recombinant inbred line (RIL) population carrying a combination stg QTL were evaluated under three environments in four replications. The results revealed that stg QTL have variable effects on severity of stalk rot diseases. Genotypes carrying either stg1 or stg3, or their combination stg1+3 expressed high levels of resistance to M. phaseolina. But, resistance to F. thapsinum required a combination of stg1 and stg3. Other stg QTL blocks, such as stg2 and stg4, did not have an effect on stalk rot resistance caused by either pathogen. Furthermore, stalk rot resistance had no significant association with levels of dhurrin and soluble sugars in the leaves.

Cereals including wheat and barley are of primary importance to ensure food security for the 21st century. A combination of lab- and field-based approaches has led to a considerably improved understanding of the importance of organ and particularly of whole-plant (monocarpic) senescence for wheat and barley yield and quality. A delicate balance between senescence timing, grain nutrient content, nutrient-use efficiency, and yield needs to be considered to (further) improve cereal varieties for a given environment and end use. The recent characterization of the Gpc-1 (NAM-1) genes in wheat and barley demonstrates the interdependence of these traits. Lines or varieties with functional Gpc-1 genes demonstrate earlier senescence and enhanced grain protein and micronutrient content but, depending on the environment, somewhat reduced yields. A major effort is needed to dissect regulatory networks centred on additional wheat and barley transcription factors and signalling pathways influencing the senescence process. Similarly, while important molecular details of nutrient (particularly nitrogen) remobilization from senescing organs to developing grains have been identified, important knowledge gaps remain. The genes coding for the major proteases involved in senescence-associated plastidial protein degradation are largely unknown. Membrane transport proteins involved in the different transport steps occurring between senescing organ (such as leaf mesophyll) cells and protein bodies in the endosperm of developing grains remain to be identified or further characterized. Existing data suggest that an improved understanding of all these steps will reveal additional, important targets for continued cereal improvement. © The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.

A stay-green phenotype enables crops to retain green leaves longer after anthesis compared with senescent types, potentially improving yield. Measuring the normalized difference vegetative index (NDVI) during the whole senescence period allows quantification of component stay-green traits contributing to a stay-green phenotype. These objective and standardized traits can be compared across genotypes and environments. Traits examined include maximum NDVI near anthesis (Nmax), senescence rate (SR), a trait integrating senescence (SGint), plus time from anthesis to onset (OnS), mid-point (MidS), and near completion (EndS) of senescence. The correlation between stay-green traits and yield was studied in eight contrasting environments ranging from well watered to severely water limited. Environments were each classified into one of the four major drought environment types (ETs) previously identified for the Australian wheat cropping system. SGint, OnS, and MidS tended to have higher values in higher yielding environments for a given genotype, as well as for higher yielding genotypes within a given environment. Correlation between specific stay-green traits and yield varied with ET. In the studied population, SGint, OnS, and MidS strongly correlated with yield in three of the four ETs which included well-watered environments (0.43-0.86), but less so in environments with only moderate water-stress after anthesis (-0.03 to 0.31). In contrast, Nmax was most highly correlated with yield under moderate post-anthesis water stress (0.31-0.43). Selection for particular stay-green traits, combinations of traits, and/or molecular markers associated with the traits could enhance genetic progress toward stay-green wheats with higher, more stable yield in both well-watered and water-limited conditions.© The Author 2016. Published by Oxford University Press on behalf of the Society for Experimental Biology.

Wheat is highly prone to terminal heat stress (HS) under late-sown conditions. Delayed-sowing is one of the preferred methods to screen the genotypes for thermotolerance under open field conditions. We investigated the effect of terminal HS on the thermotolerance of four popular genotypes of wheat i.e. WR544, HD2967, HD2932, and HD2285 under field condition. We observed significant variations in the biochemical parameters like protein content, antioxidant activity, proline and total reducing sugar content in leaf, stem, and spike under normal (26 +/- 2 degrees C) and terminal HS (36 +/- 2 degrees C) conditions. Maximum protein, sugars and proline was observed in HD2967, as compared to other cultivars under terminal HS. Wheat cv. HD2967 showed more adaptability to the terminal HS. Differential protein-profiling in leaves, stem and spike of HD2967 under normal (26 +/- 2 degrees C) and terminal HS (36 +/- 2 degrees C) showed expression of some unique protein spots. MALDI-TOF/MS analysis showed the DEPs as RuBisCO (Rub), RuBisCO activase (Rca), oxygen evolving enhancer protein (OEEP), hypothetical proteins, etc. Expression analysis of genes associated with photosynthesis (Rub and Rca) and starch biosynthesis pathway (AGPase, SSS and SBE) showed significant variations in the expression under terminal HS. HD2967 showed better performance, as compared to other cultivars under terminal HS. SSS activity observed in HD2967 showed more stability under terminal HS, as compared with other cultivars. Triggering of different biochemical parameters in response to terminal HS was observed to modulate the plasticity of carbon assimilatory pathway. The identified DEPs will enrich the proteomic resources of wheat and will provide a potential biochemical marker for screening wheat germplasm for thermotolerance. The model hypothesized will help the researchers to work in a more focused way to develop terminal heat tolerant wheat without compromising with the quality and quantity of grains.

Enhancing the nutritional value of food crops is a means of improving human nutrition and health. We report here the positional cloning of Gpc-B1, a wheat quantitative trait locus associated with increased grain protein, zinc, and iron content. The ancestral wild wheat allele encodes a NAC transcription factor (NAM-B1) that accelerates senescence and increases nutrient remobilization from leaves to developing grains, whereas modern wheat varieties carry a nonfunctional NAM-B1 allele. Reduction in RNA levels of the multiple NAM homologs by RNA interference delayed senescence by more than 3 weeks and reduced wheat grain protein, zinc, and iron content by more than 30%.

In this review, we present a comprehensive revisit of past research and advances developed on the stay-green (SG) paradigm. The study aims to provide an application-focused review of the SG phenotypes as crop residuals for bioenergy. Little is known about the SG trait as a germplasm enhancer resource for energy storage as a system for alternative energy. Initially described as a single locus recessive trait, SG was shortly after reported as a quantitative trait governed by complex physiological and metabolic networks including chlorophyll efficiency, nitrogen contents, nutrient remobilization and source-sink balance. Together with the fact that phenotyping efforts have improved rapidly in the last decade, new approaches based on sensing technologies have had an impact in SG identification. Since SG is linked to delayed senescence, we present a review of the term senescence applied to crop residuals and bioenergy. Firstly, we discuss the idiosyncrasy of senescence. Secondly, we present biological processes that determine the fate of senescence. Thirdly, we present the genetics underlying SG for crop-trait improvement in different crops. Further, this review explores the potential uses of senescence for bioenergy crops. Finally, we discuss how high-throughput phenotyping methods assist new technologies such as genomic selection in a cost-efficient manner.

<div style="line-height: 150%">Abstract <br>Stay green (SG) maize was found to have higher grain yield and post-silking nitrogen (N) uptake (PostN) compared with a non-stay green (NSG) hybrid.&nbsp; To understand the effects of plant density on grain yield (GY) and N efficiency in modern maize hybrids, we compared two modern hybrids (SG hybrid DY508 and NSG hybrid NH101) with similar maturity ratings at three plant densities (45 000, 60 000, and 75 000 pl ha^&ndash;1) in 2014 and 2015.&nbsp; GY, leaf senescence, dry matter (DM) accumulation, N accumulation, PostN, and post-silking N remobilization (RemN) were analyzed.&nbsp; DY508 and NH101 had similar GY, but DY508 had higher thousand kernel weight (TKW) and lower kernel number (KN) than NH101.&nbsp; Plant density significantly increased GY in the two hybrids.&nbsp; On average, over the two years, plant density improved GY in DY508 and NH101 by 18.5 and 11.1%, respectively, but there were no differences in total dry matter (TDM) and post-silking DM (PostDM) between the two hybrids.&nbsp; Plant density improved leaf N, stem N, and grain N at the silking and maturity stages in 2014 and 2015.&nbsp; DY508 was lower in harvest index (HI), nitrogen harvest index (NHI), and grain N concentration (GNC) than NH101.&nbsp; Grain N in DY508 was 2.61 kg ha^&ndash;1 less than in NH101, and this was caused by lower GNC and leaf RemN.&nbsp; On the average, DY508 was 1.62 kg ha^&ndash;1 less in leaf remobilized N (leaf RemN) than NH101, but was similar in stem remobilized N (stem RemN; 2.47 kg ha^&ndash;1 vs. 3.41 kg ha^&ndash;1).&nbsp; Maize hybrid DY508 shows delayed leaf senescence in the upper and bottom canopy layers in the later stages of growth.&nbsp; The present study provides evidence that the NH101, which has rapid leaf senescence at the late grain-filling stage, has gained equivalent GY and higher leaf RemN, and was more efficient in N utilization.</div>

Senescence is a highly organized and well-regulated process. As much as 75% of total cellular nitrogen may be located in mesophyll chloroplasts of C(3)-plants. Proteolysis of chloroplast proteins begins in an early phase of senescence and the liberated amino acids can be exported to growing parts of the plant (e.g. maturing fruits). Rubisco and other stromal enzymes can be degraded in isolated chloroplasts, implying the involvement of plastidial peptide hydrolases. Whether or not ATP is required and if stromal proteins are modified (e.g. by reactive oxygen species) prior to their degradation are questions still under debate. Several proteins, in particular cysteine proteases, have been demonstrated to be specifically expressed during senescence. Their contribution to the general degradation of chloroplast proteins is unclear. The accumulation in intact cells of peptide fragments and inhibitor studies suggest that multiple degradation pathways may exist for stromal proteins and that vacuolar endopeptidases might also be involved under certain conditions. The breakdown of chlorophyll-binding proteins associated with the thylakoid membrane is less well investigated. The degradation of these proteins requires the simultaneous catabolism of chlorophylls. The breakdown of chlorophylls has been elucidated during the last decade. Interestingly, nitrogen present in chlorophyll is not exported from senescencing leaves, but remains within the cells in the form of linear tetrapyrrolic catabolites that accumulate in the vacuole. The degradation pathways for chlorophylls and chloroplast proteins are partially interconnected.