Acclimation of leaf photosynthesis and respiration to warming in field-grown wheat.

Climate change and future warming will significantly affect crop yield. The capacity of crops to dynamically adjust physiological processes (i.e., acclimate) to warming might improve overall performance. Understanding and quantifying the degree of acclimation in field crops could ensure better parameterization of crop and Earth System models and predictions of crop performance. We hypothesized that for field-grown wheat, when measured at a common temperature (25°C), crops grown under warmer conditions would exhibit acclimation, leading to enhanced crop performance and yield. Acclimation was defined as (a) decreased rates of net photosynthesis at 25°C (A25 ) coupled with lower maximum carboxylation capacity (Vcmax25 ), (b) reduced leaf dark respiration at 25°C (both in terms of O2 consumption Rdark _O225 and CO2 efflux Rdark _CO225 ) and (c) lower Rdark _CO225 to Vcmax25 ratio. Field experiments were conducted over two seasons with 20 wheat genotypes, sown at three different planting dates, to test these hypotheses. Leaf-level CO2 -based traits (A25 , Rdark _CO225 and Vcmax25 ) did not show the classic acclimation responses that we hypothesized; by contrast, the hypothesized changes in Rdark_ O2 were observed. These findings have implications for predictive crop models that assume similar temperature response among these physiological processes and for predictions of crop performance in a future warmer world.

Effect of gibberellin-sensitive Rht18 and gibberellin-insensitive Rht-D1b dwarfing genes on vegetative and reproductive growth in bread wheat.

Gibberellin (GA)-insensitive dwarfing genes Rht-B1b and Rht-D1b that are responsible for the 'Green Revolution' have been remarkably successful in wheat improvement globally. However, these alleles result in shorter coleoptiles and reduced vigour, and hence poor establishment and growth in some environments. Rht18, on the other hand, is a GA-sensitive, dominant gene with potential to overcome some of the early growth limitations associated with Rht-B1b and Rht-D1b. We assessed progeny from both a biparental and a backcross population that contained tall, single dwarf, and double dwarf lines, to determine whether Rht18 differs from Rht-D1b and hence verify its value in wheat improvement. Progeny with Rht18 had an almost identical height to lines with Rht-D1b, and both were ~26% shorter than the tall lines, with the double dwarf 13% shorter again. However, coleoptile length of Rht18 was 42% longer than that of Rht-D1b. We detected no differences in time to terminal spikelet and anthesis, and few differences in stem or spike growth. Both dwarfing genes diverted more dry matter to the spike than tall lines from prior to heading. No differences were detected between Rht18 and Rht-D1b that could prevent the adoption of Rht18 in wheat breeding to overcome some of the limitations associated with the 'Green Revolution' genes.

Wide variation in the suboptimal distribution of photosynthetic capacity in relation to light across genotypes of wheat.

Suboptimal distribution of photosynthetic capacity in relation to light among leaves reduces potential whole-canopy photosynthesis. We quantified the degree of suboptimality in 160 genotypes of wheat by directly measuring photosynthetic capacity and daily irradiance in flag and penultimate leaves. Capacity per unit daily irradiance was systematically lower in flag than penultimate leaves in most genotypes, but the ratio (γ) of capacity per unit irradiance between flag and penultimate leaves varied widely across genotypes, from less than 0.5 to over 1.2. Variation in γ was most strongly associated with differences in photosynthetic capacity in penultimate leaves, rather than with flag leaf photosynthesis or canopy light penetration. Preliminary genome-wide association analysis identified nine strong marker-trait associations with this trait, which should be validated in future work in other environments and/or materials. Our modelling suggests canopy photosynthesis could be increased by up to 5 % under sunny conditions by harnessing this variation through selective breeding for increased γ.

Deeper roots associated with cooler canopies, higher normalized difference vegetation index, and greater yield in three wheat populations grown on stored soil water.

Simple and repeatable methods are needed to select for deep roots under field conditions. A large-scale field experiment was conducted to assess the association between canopy temperature (CT) measured by airborne thermography and rooting depth determined by the core-break method. Three wheat populations, C306×Westonia (CW), Hartog×Drysdale (HD), and Sundor×Songlen (SS), were grown on stored soil water in NSW Australia in 2017 (n=196-252). Cool and warm CT extremes ('tails') were cored after harvest (13-32% of each population). Rooting depth was significantly correlated with CT at late flowering (r= -0.25, -0.52, and -0.23 for CW, HD, and SS, respectively, P<0.05 hereafter), with normalized difference vegetation index (NDVI) at early grain filling (r=0.30-0.39), and with canopy height (r=0.23-0.48). The cool tails showed significantly deeper roots than the respective warm tails by 8.1 cm and 6.2 cm in CW and HD, and correspondingly, greater yields by an average 19% and 7%, respectively. This study highlighted that CT measured rapidly by airborne thermography or NDVI at early grain filling could be used to guide selection of lines with deeper roots to increase wheat yields. The remote measurement methods in this study were repeatable and high throughput, making them well suited to use in breeding programmes.

Exploring high temperature responses of photosynthesis and respiration to improve heat tolerance in wheat.

High temperatures account for major wheat yield losses annually and, as the climate continues to warm, these losses will probably increase. Both photosynthesis and respiration are the main determinants of carbon balance and growth in wheat, and both are sensitive to high temperature. Wheat is able to acclimate photosynthesis and respiration to high temperature, and thus reduce the negative affects on growth. The capacity to adjust these processes to better suit warmer conditions stands as a potential avenue toward reducing heat-induced yield losses in the future. However, much remains to be learnt about such phenomena. Here, we review what is known of high temperature tolerance in wheat, focusing predominantly on the high temperature responses of photosynthesis and respiration. We also identify the many unknowns that surround this area, particularly with respect to the high temperature response of wheat respiration and the consequences of this for growth and yield. It is concluded that further investigation into the response of photosynthesis and respiration to high temperature could present several methods of improving wheat high temperature tolerance. Extending our knowledge in this area could also lead to more immediate benefits, such as the enhancement of current crop models.

Rate of photosynthetic induction in fluctuating light varies widely among genotypes of wheat.

Crop photosynthesis and yield are limited by slow photosynthetic induction in sunflecks. We quantified variation in induction kinetics across diverse genotypes of wheat for the first time. Following a preliminary study that hinted at wide variation in induction kinetics across 58 genotypes, we grew 10 genotypes with contrasting responses in a controlled environment and quantified induction kinetics of carboxylation capacity (Vcmax) from dynamic A versus ci curves after a shift from low to high light (from 50 µmol m-2 s-1 to 1500 µmol m-2 s-1), in five flag leaves per genotype. Within-genotype median time for 95% induction (t95) of Vcmax varied 1.8-fold, from 5.2 min to 9.5 min. Our simulations suggest that non-instantaneous induction reduces daily net carbon gain by up to 15%, and that breeding to speed up Vcmax induction in the slowest of our 10 genotypes to match that in the fastest genotype could increase daily net carbon gain by up to 3.4%, particularly for leaves in mid-canopy positions (cumulative leaf area index ≤1.5 m2 m-2), those that experience predominantly short-duration sunflecks, and those with high photosynthetic capacities.

Wheats developed for high yield on stored soil moisture have deep vigorous root systems.

Many rainfed wheat production systems are reliant on stored soil water for some or all of their water inputs. Selection and breeding for root traits could result in a yield benefit; however, breeding for root traits has traditionally been avoided due to the difficulty of phenotyping mature root systems, limited understanding of root system development and function, and the strong influence of environmental conditions on the phenotype of the mature root system. This paper outlines an international field selection program for beneficial root traits at maturity using soil coring in India and Australia. In the rainfed areas of India, wheat is sown at the end of the monsoon into hot soils with a quickly receding soil water profile; in season water inputs are minimal. We hypothesised that wheat selected and bred for high yield under these conditions would have deep, vigorous root systems, allowing them to access and utilise the stored soil water at depth around anthesis and grain-filling when surface layers were dry. The Indian trials resulted in 49 lines being sent to Australia for phenotyping. These lines were ranked against 41 high yielding Australian lines. Variation was observed for deep root traits e.g. in eastern Australia in 2012, maximum depth ranged from 118.8 to 146.3cm. There was significant variation for root traits between sites and years, however, several Indian genotypes were identified that consistently ranked highly across sites and years for deep rooting traits.

Physiological perspectives of reduced tillering and stunting in the tiller inhibition (tin) mutant of wheat.

The number of tillers established in cereal crops far exceeds the number that end up being grain bearing at maturity. Improving the economy in tillering has been proposed to improve cereal yields in both favourable and unfavourable environments. The tiller inhibition mutant (tin) is potentially useful for breeding varieties with a greater economy of tillering. However, its tendency to stunting under long day and low temperatures has limited its use. Recently, the inhibition of tillering in tin has been linked to precocious development of solid basal internodes that compete for sucrose and possibly other resources with the growing tiller buds leading to their developmental arrest. Although the physiological basis of stunting in tin is unknown, both inhibition of tillering and stunting begin during the transition from vegetative to reproductive phase indicating a common physiological basis for both. In this review, we provide overall perspectives for the physiological basis of tiller inhibition and stunting in tin and suggest the direction of research in the future.

Inhibition of tiller bud outgrowth in the tin mutant of wheat is associated with precocious internode development.

Tillering (branching) is a major yield component and, therefore, a target for improving the yield of crops. However, tillering is regulated by complex interactions of endogenous and environmental signals, and the knowledge required to achieve optimal tiller number through genetic and agronomic means is still lacking. Regulatory mechanisms may be revealed through physiological and molecular characterization of naturally occurring and induced tillering mutants in the major crops. Here we characterize a reduced tillering (tin, for tiller inhibition) mutant of wheat (Triticum aestivum). The reduced tillering in tin is due to early cessation of tiller bud outgrowth during the transition of the shoot apex from the vegetative to the reproductive stage. There was no observed difference in the development of the main stem shoot apex between tin and the wild type. However, tin initiated internode development earlier and, unlike the wild type, the basal internodes in tin were solid rather than hollow. We hypothesize that tin represents a novel type of reduced tillering mutant associated with precocious internode elongation that diverts sucrose (Suc) away from developing tillers. Consistent with this hypothesis, we have observed upregulation of a gene induced by Suc starvation, downregulation of a Suc-inducible gene, and a reduced Suc content in dormant tin buds. The increased expression of the wheat Dormancy-associated (DRM1-like) and Teosinte Branched1 (TB1-like) genes and the reduced expression of cell cycle genes also indicate bud dormancy in tin. These results highlight the significance of Suc in shoot branching and the possibility of optimizing tillering by manipulating the timing of internode elongation.

Genomic regions for canopy temperature and their genetic association with stomatal conductance and grain yield in wheat.

Stomata are the site of CO2 exchange for water in a leaf. Variation in stomatal control offers promise in genetic improvement of transpiration and photosynthetic rates to improve wheat performance. However, techniques for estimating stomatal conductance (SC) are slow, limiting potential for efficient measurement and genetic modification of this trait. Genotypic variation in canopy temperature (CT) and leaf porosity (LP), as surrogates for SC, were assessed in three wheat mapping populations grown under well-watered conditions. The range and resulting genetic variance were large but not always repeatable across days and years for CT and LP alike. Leaf-to-leaf variation was large for LP, reducing heritability to near zero on a single-leaf basis. Replication across dates and years increased line-mean heritability to ~75% for both CT and LP. Across sampling dates and populations, CT showed a large, additive genetic correlation with LP (rg=-0.67 to -0.83) as expected. Genetic increases in pre-flowering CT were associated with reduced final plant height and both increased harvest index and grain yield but were uncorrelated with aerial biomass. In contrast, post-flowering, cooler canopies were associated with greater aerial biomass and increased grain number and yield. A multi-environment QTL analysis identified up to 16 and 15 genomic regions for CT and LP, respectively, across all three populations. Several of the LP and CT QTL co-located with known QTL for plant height and phenological development and intervals for many of the CT and LP quantitative trait loci (QTL) overlapped, supporting a common genetic basis for the two traits. Notably, both Rht-B1b and Rht-D1b dwarfing alleles were paradoxically positive for LP and CT (i.e. semi-dwarfs had higher stomatal conductance but warmer canopies) highlighting the issue of translation from leaf to canopy in screening for greater transpiration. The strong requirement for repeated assessment of SC suggests the more rapid CT assessment may be of greater value for indirect screening of high or low SC among large numbers of early-generation breeding lines. However, account must be taken of variation in development and canopy architecture when interpreting performance and selecting breeding lines on the basis of CT.

Genetic control of duration of pre-anthesis phases in wheat (Triticum aestivum L.) and relationships to leaf appearance, tillering, and dry matter accumulation.

The duration of pre-anthesis developmental phases is of interest in breeding for improved adaptation and yield potential in temperate cereals. Yet despite numerous studies on the genetic control of anthesis (flowering) time and floral initiation, little is known about the genetic control of other pre-anthesis phases. Furthermore, little is known about the effect that changes in the duration of pre-anthesis phases could have on traits related to leaf appearance and tillering, or dry matter accumulation before terminal spikelet initiation (TS). The genetic control of the leaf and spikelet initiation phase (LS; from sowing to TS), the stem elongation phase (SE; from TS to anthesis), and, within the latter, from TS to flag leaf appearance and from then to anthesis, was studied in two doubled-haploid, mapping bread wheat populations, Cranbrook × Halberd and CD87 × Katepwa, in two field experiments (ACT and NSW, Australia). The lengths of phases were estimated from measurements of both TS and the onset of stem elongation. Dry weight per plant before TS, rate of leaf appearance, tillering rate, maximum number of tillers and number of leaves, and dry weight per plant at TS were also estimated in the Cranbrook × Halberd population. More genomic regions were identified for the length of the different pre-anthesis phases than for total time to anthesis. Although overall genetic correlations between LS and SE were significant and positive, independent genetic variability between LS and SE, and several quantitative trait loci (QTLs) with different effects on both phases were found in the two populations. Several of these QTLs (which did not seem to coincide with reported major genes) could be of interest for breeding purposes since they were only significant for either LS or SE. There was no relationship between LS and the rate of leaf appearance. LS was strongly and positively correlated with dry weight at TS but only slightly negatively correlated with early vigour (dry weight before TS). Despite significant genetic correlations between LS and some tillering traits, shortening LS so as to lengthen SE without modifying total time to anthesis would not necessarily reduce tillering capacity, as QTLs for tillering traits did not coincide with those QTLs significant only for LS or SE. Therefore, the study of different pre-anthesis phases is relevant for a better understanding of genetic factors regulating developmental time and may offer new tools for fine-tuning it in breeding for both adaptability and yield potential.

Abiotic stress and control of grain number in cereals.

Grain number is the only yield component that is directly associated with increased grain yield in important cereal crops like wheat. Historical yield studies show that increases in grain yield are always accompanied by an increase in grain number. Adverse weather conditions can cause severe fluctuations in grain yield and substantial yield losses in cereal crops. The problem is global and despite its impact on world food production breeding and selection approaches have only met with limited success. A specific period during early reproductive development, the young microspore stage of pollen development, is extremely vulnerable to abiotic stress in self-fertilising cereals (wheat, rice, barley, sorghum). A better understanding of the physiological and molecular processes that lead to stress-induced pollen abortion may provide us with the key to finding solutions for maintaining grain number under abiotic stress conditions. Due to the complexity of the problem, stress-proofing our main cereal crops will be a challenging task and will require joint input from different research disciplines.

Importance of pre-anthesis anther sink strength for maintenance of grain number during reproductive stage water stress in wheat.

Reproductive stage water stress leads to spikelet sterility in wheat. Whereas drought stress at anthesis affects mainly grain size, stress at the young microspore stage of pollen development is characterized by abortion of pollen development and reduction in grain number. We identified genetic variability for drought tolerance at the reproductive stage. Drought-tolerant wheat germplasm is able to maintain carbohydrate accumulation in the reproductive organs throughout the stress treatment. Starch depletion in the ovary of drought-sensitive wheat is reversible upon re-watering and cross-pollination experiments indicate that the ovary is more resilient than the anther. The effect on anthers and pollen fertility is irreversible, suggesting that pollen sterility is the main cause of grain loss during drought conditions in wheat. The difference in storage carbohydrate accumulation in drought-sensitive and drought-tolerant wheat is correlated with differences in sugar profiles, cell wall invertase gene expression and expression of fructan biosynthesis genes in anther and ovary (sucrose : sucrose 1-fructosyl-transferase, 1-SST; sucrose : fructan 6-fructosyl-transferase, 6-SFT). Our results indicate that the ability to control and maintain sink strength and carbohydrate supply to anthers may be the key to maintaining pollen fertility and grain number in wheat and this mechanism may also provide protection against other abiotic stresses.

Genotypic variation in water-soluble carbohydrate accumulation in wheat.

The water-soluble carbohydrate (WSC) that accumulates in the stems of wheat during growth can be an important contributor to grain filling, particularly under conditions when assimilation is limited, such as during end-of-season drought. WSC concentration was measured at anthesis across a diverse set of wheat genotypes over multiple environments. Environmental differences in WSC concentration were large (means for the set ranging between 108 and 203 mg g-1 dry weight), and there were significant and repeatable differences in WSC accumulation among genotypes (means ranging from 112 to 213 mg g-1 dry weight averaged across environments), associated with large broad-sense heritability (H = 0.90 ± 0.12). These results suggest that breeding for high WSC should be possible in wheat. The composition of the WSC, examined in selected genotypes, indicated that the variation in total WSC was attributed mainly to variation in the fructan component, with the other major soluble carbohydrates, sucrose and hexose, varying less. The degree of polymerisation (DP) of fructo-oligosaccharides was up to ~13 in samples where higher levels of WSC were accumulated, owing either to genotype or environment, but the higher DP components (DP > 6) were decreased in samples of lower total WSC. The results are consistent with fructan biosynthesis occurring via a sequential mechanism that is dependent on the availability of sucrose, and differences in WSC contents of genotypes are unlikely to be due to major mechanistic differences.

Influence of the gibberellin-sensitive Rht8 dwarfing gene on leaf epidermal cell dimensions and early vigour in wheat (Triticum aestivum L.).

BACKGROUND AND AIMS: The gibberellin-insensitive Rht-B1b and Rht-D1b dwarfing genes are known to reduce the size of cells in culms, leaves and coleoptiles of wheat. Resulting leaf area development of gibberellin-insensitive wheats is poor compared to standard height (Rht-B1a and Rht-D1a) genotypes. Alternative dwarfing genes to Rht-B1b and Rht-D1b are available that reduce plant height, such as the gibberellin-responsive Rht8 gene. This study aims to investigate if Rht8 has a similar dwarfing effect on the size of leaf cells to reduce leaf area. METHODS: The effect of Rht8 on cell size and leaf area was assessed in four types of epidermal cells (interstomatal, long, sister and bulliform) measured on leaf 2 of standard height (rht8) and semi-dwarf (Rht8) doubled-haploid lines (DHLs). The DHLs were derived from a cross between very vigorous, standard height (rht8) ('Vigour18') and less vigorous, semi-dwarf (Rht8) ('Chuan-Mai 18') parents. KEY RESULTS: Large differences were observed in seedling vigour between the parents, where 'Vigour18' had a much greater plant leaf area than 'Chuan-Mai 18'. Accordingly, 'Vigour18' had on average longer, wider and more epidermal cells and cell files than 'Chuan-Mai 18'. Although there was correspondingly large genotypic variation among DHLs for these traits, the contrast between semi-dwarf Rht8 and tall rht8 DHLs revealed no difference in the size of leaf 2 or average cell characteristics. Hence, these traits were independent of plant height and therefore Rht8 in the DHLs. Correlations for leaf and average cell size across DHLs revealed a strong and positive relationship between leaf width and cell files, while the relationships between leaf and cell width, and leaf and cell length were not statistically different. The relative contribution of the four cell types (long, sister, interstomatal and bulliform) to leaf size in the parents, comparative controls and DHLs is discussed. CONCLUSIONS: Despite a large range in early vigour among the DHLs, none of the DHLs attained the leaf area or epidermal cell size and numbers of the vigorous rht8 parent. Nonetheless, the potential exists to increase the early vigour of semi-dwarf wheats by using GA-sensitive dwarfing genes such as Rht8.

The effect of different height reducing genes on the early growth of wheat.

Genes that reduce height without compromising seedling vigour or coleoptile length have great potential for wheat improvement. We therefore investigated the effects of various reduced height (Rht) genes on the early stages of plant development, using a combination of near isogenic, recombinant, mutant and wild type comparisons. Gibberellin (GA) insensitivity caused by Rht-B1b or Rht-D1b was associated with reduced leaf elongation rate and coleoptile length. Similar results were found for two other sources of dwarfing, Rht11 and Rht17. We found one class of Rht genes (e.g. Rht8) which had no effect on coleoptile length, leaf elongation rate or responsiveness to GA, indicating that these dwarfing genes may act later in wheat development to reduce height and increase harvest index, without affecting early growth. A third class of Rht genes was found in three durum backgrounds. These had reduced coleoptile lengths and leaf elongation rates, but had a greater response to GA than the corresponding tall varieties. We discuss these results in relation to the possible mechanisms underlying the reduction in height and the suitability of the different Rht genes for wheat improvement.

Environmental effects on stunting and the expression of a tiller inhibition (tin) gene in wheat.

A recessive gene (tin) that inhibits tillering in wheat (Triticum aestivum L.), and that may be important in the redirection of assimilate from unproductive to productive tillers, has been reported. However, this gene has also been associated with a fatal condition known as 'stunting'. The severity of this phenomenon has been shown to increase when plants are grown under long photoperiods and at low temperatures. The objectives of this study were to observe how the expression of the tin gene varied in different genetic backgrounds, in addition to obtaining a better understanding of environmental factors that may affect both tillering and stunting in lines with the tin gene. Plants were grown outdoors in Canberra, Australia, at various times throughout the year, as well as under controlled conditions where photoperiod, temperature and light intensity were varied. The inhibition of tillers resulting from the presence of the tin gene was most extreme in summer, autumn and spring (up to 90% reduction in tillering). However, when sown in late autumn and winter, tillering was reduced by between 30-50% for the tin lines compared with their near-isogenic parents. Reduced tillering in the tin lines was due to an earlier cessation of tillering rather than a reduced rate. Stunting was frequently observed in some lines more than others when plants were grown under long days and when temperatures were low. The daily minimum temperature, rather than the average daily temperature, was associated with stunting. The duration of the dark period also influenced stunting, with a longer dark period reducing the incidence of stunting from almost 100% to 0%. In all experiments where irradiance was increased, stunting also increased. In addition, elevated CO2 also increased growth characteristics associated with stunting. It is concluded that stunting is associated with a high assimilate supply to the main stem shoot apex before the time of floral initiation. This is caused by an inhibition of tillering and a high photothermal quotient. Leaf length was found to be a good indicator of stunting severity, with stunted plants producing shorter leaves than those plants which failed to stunt. Measurements of leaf length indicated that stunting is induced when the second leaf is expanding.