Physiological adaptation to irradiance in duckweeds is species and accession specific and depends on light habitat niche.

Duckweeds span 36 species of free-floating aquatic organisms with body sizes ranging from 2 mm to 10 mm, where each plant body plan is reduced to a largely leaf-like structure. As an emerging crop, their fast growth rates offer potential for cultivation in closed systems. We describe a novel UK collection derived from low light (dLL) or high light (dHL) habitats, profiled for growth, photosynthesis, and photoprotection (non-photochemical quenching, NPQ) responses. Twenty-three accessions of three Lemna species and one Spirodela polyrhiza were grown under relatively low light (LL: 100 µmol m-2 s-1) and high light (HL: 350 µmol m-2 s-1) intensities. We observed broad within- and between-species level variation in photosynthesis acclimation. Duckweeds grown under HL exhibited a lower growth rate, biomass, chlorophyll, and quantum yield of photosynthesis. In HL compared with LL, carotenoid de-epoxidation state and NPQ were higher, whilst PSII efficiency (φPSII) and Chl a:b ratios were unchanged. The dLL plants showed relatively stronger acclimation to HL compared with dHL plants, especially Lemna japonica accessions. These achieved faster growth in HL with concurrent higher carotenoid levels and NPQ, and less degradation of chlorophyll. We conclude that these data support local adaptation to the light environment in duckweed affecting acclimation in controlled conditions.

Night-time warming in the field reduces nocturnal stomatal conductance and grain yield but does not alter daytime physiological responses.

Global nocturnal temperatures are rising more rapidly than daytime temperatures and have a large effect on crop productivity. In particular, stomatal conductance at night (gsn ) is surprisingly poorly understood and has not been investigated despite constituting a significant proportion of overall canopy water loss. Here, we present the results of 3 yr of field data using 12 spring Triticum aestivum genotypes which were grown in NW Mexico and subjected to an artificial increase in night-time temperatures of 2°C. Under nocturnal heating, grain yields decreased (1.9% per 1°C) without significant changes in daytime leaf-level physiological responses. Under warmer nights, there were significant differences in the magnitude and decrease in gsn , values of which were between 9 and 33% of daytime rates while respiration appeared to acclimate to higher temperatures. Decreases in grain yield were genotype-specific; genotypes categorised as heat tolerant demonstrated some of the greatest declines in yield in response to warmer nights. We conclude the essential components of nocturnal heat tolerance in wheat are uncoupled from resilience to daytime temperatures, raising fundamental questions for physiological breeding. Furthermore, this study discusses key physiological traits such as pollen viability, root depth and irrigation type may also play a role in genotype-specific nocturnal heat tolerance.

A 'wiring diagram' for sink strength traits impacting wheat yield potential.

Identifying traits for improving sink strength is a bottleneck to increasing wheat yield. The interacting processes determining sink strength and yield potential are reviewed and visualized in a set of 'wiring diagrams', covering critical phases of development (and summarizing known underlying genetics). Using this framework, we reviewed and assembled the main traits determining sink strength and identified research gaps and potential hypotheses to be tested for achieving gains in sink strength. In pre-anthesis, grain number could be increased through: (i) enhanced spike growth associated with optimized floret development and/or a reduction in specific stem-internode lengths and (ii) improved fruiting efficiency through an accelerated rate of floret development, improved partitioning between spikes, or optimized spike cytokinin levels. In post-anthesis, grain, sink strength could be augmented through manipulation of grain size potential via ovary size and/or endosperm cell division and expansion. Prospects for improving spike vascular architecture to support all rapidly growing florets, enabling the improved flow of assimilate, are also discussed. Finally, we considered the prospects for enhancing grain weight realization in relation to genetic variation in stay-green traits as well as stem carbohydrate remobilization. The wiring diagrams provide a potential workspace for breeders and crop scientists to achieve yield gains in wheat and other field crops.

A 'wiring diagram' for source strength traits impacting wheat yield potential.

Source traits are currently of great interest for the enhancement of yield potential; for example, much effort is being expended to find ways of modifying photosynthesis. However, photosynthesis is but one component of crop regulation, so sink activities and the coordination of diverse processes throughout the crop must be considered in an integrated, systems approach. A set of 'wiring diagrams' has been devised as a visual tool to integrate the interactions of component processes at different stages of wheat development. They enable the roles of chloroplast, leaf, and whole-canopy processes to be seen in the context of sink development and crop growth as a whole. In this review, we dissect source traits both anatomically (foliar and non-foliar) and temporally (pre- and post-anthesis), and consider the evidence for their regulation at local and whole-plant/crop levels. We consider how the formation of a canopy creates challenges (self-occlusion) and opportunities (dynamic photosynthesis) for components of photosynthesis. Lastly, we discuss the regulation of source activity by feedback regulation. The review is written in the framework of the wiring diagrams which, as integrated descriptors of traits underpinning grain yield, are designed to provide a potential workspace for breeders and other crop scientists that, along with high-throughput and precision phenotyping data, genetics, and bioinformatics, will help build future dynamic models of trait and gene interactions to achieve yield gains in wheat and other field crops.

Chlorophyll fluorescence-based high-throughput phenotyping facilitates the genetic dissection of photosynthetic heat tolerance in African (Oryza glaberrima) and Asian (Oryza sativa) rice.

Rising temperatures and extreme heat events threaten rice production. Half of the global population relies on rice for basic nutrition, and therefore developing heat-tolerant rice is essential. During vegetative development, reduced photosynthetic rates can limit growth and the capacity to store soluble carbohydrates. The photosystem II (PSII) complex is a particularly heat-labile component of photosynthesis. We have developed a high-throughput chlorophyll fluorescence-based screen for photosynthetic heat tolerance capable of screening hundreds of plants daily. Through measuring the response of maximum PSII efficiency to increasing temperature, this platform generates data for modelling the PSII-temperature relationship in large populations in a small amount of time. Coefficients from these models (photosynthetic heat tolerance traits) demonstrated high heritabilities across African (Oryza glaberrima) and Asian (Oryza sativa, Bengal Assam Aus Panel) rice diversity sets, highlighting valuable genetic variation accessible for breeding. Genome-wide association studies were performed across both species for these traits, representing the first documented attempt to characterize the genetic basis of photosynthetic heat tolerance in any species to date. A total of 133 candidate genes were highlighted. These were significantly enriched with genes whose predicted roles suggested influence on PSII activity and the response to stress. We discuss the most promising candidates for improving photosynthetic heat tolerance in rice.

A multiscale mathematical model describing the growth and development of bambara groundnut.

A principal objective in agriculture is to maximise food production; this is particularly relevant with the added demands of an ever increasing population, coupled with the unpredictability that climate change brings. Further improvements in productivity can only be achieved with an increased understanding of plant and crop processes. In this respect, mathematical modelling of plants and crops plays an important role. In this paper we present a two-scale mathematical model of crop yield that accounts for plant growth and canopy interactions. A system of nonlinear ordinary differential equations (ODEs) is formulated to describe the growth of each individual plant, where equations are coupled via a term that describes plant competition via canopy-canopy interactions. A crop of greenhouse plants is then modelled via an agent based modelling approach in which the growth of each plant is described via our system of ODEs. The model is formulated for the African drought tolerant legume bambara groundnut (Vigna subterranea), which is currently being investigated as a food source in light of climate change and food insecurity challenges. Our model allows us to account for plant diversity and also investigate the effect of individual plant traits (e.g. plant canopy size and planting distance) on the yield of the overall crop. Informed with greenhouse data, model results show that plant positioning relative to other plants has a large impact on individual plant yield. Variation in physiological plant traits from genetic diversity and the environmental effects lead to experimentally observed variations in crop yield. These traits include plant height, plant carrying capacity, leaf accumulation rate and canopy spread. Of these traits plant height and ground cover growth rates are found to have the greatest impact on crop yield. We also consider a range of different planting arrangements (uniform grid, staggered grid, circular rings and random allocation) and find that the staggered grid leads to the greatest crop yield (6% more compared to uniform grid). Whilst formulated specifically for bambara groundnut, the generic formulation of our model means that with changes to certain parameter's, it may be extended to other crop species that form a canopy.

The rice EP3 and OsFBK1 E3 ligases alter plant architecture and flower development, and affect transcript accumulation of microRNA pathway genes and their targets.

ERECTA PANICLE 3 (EP3) and ORYZA SATIVA F-BOX KELCH 1 (OsFBK1) proteins share 57% and 54% sequence identity with the Arabidopsis F-box protein HAWAIIAN SKIRT (HWS). Previously we showed that EP3 is a functional orthologue of HWS. Here we demonstrate that OsFBK1 is another functional orthologue of HWS and show the complexity of interaction between EP3 and OsFBK1 genes at different developmental stages of the plant. qRT-PCR expression analyses and studies of EP3-GFP and OsFBK1-RFP promoter reporter lines demonstrate that although EP3 and OsFBK1 expression can be detected in the same tissues some cells exclusively express EP3 or OsFBK1 whilst others co-express both genes. Loss, reduction or gain-of-function lines for EP3 and OsFBK1, show that EP3 and OsFBK1 affect plant architecture, organ size, floral organ number and size, floral morphology, pollen viability, grain size and weight. We have identified the putative orthologue genes of the rice microRNA pathway for ORYZA SATIVA DAWDLE (OsDDL) and ORYZA SATIVA SERRATE (OsSE), and demonstrated that EP3 and OsFBK1 affect their transcript levels as well as those of CROWN ROOT DEFECT 1/ORYZA SATIVA Exportin-5 HASTY (CRD1/OsHST), ORYZA SATIVA DICER-LIKE 1 (OsDCL) and ORYZA SATIVA WEAVY LEAF1 (OsWAF1). We show that EP3 affects OsPri-MIR164, OsNAM1 and OsNAC1 transcript levels. OsNAC1 transcripts are modified by OsFBK1, suggesting two independent regulatory pathways, one via EP3 and OsMIR164 and the other via OsFBK1. Our data propose that EP3 and OsFBK1 conjointly play similar roles in rice to how HWS does in Arabidopsis.

Sunflecks in the upper canopy: dynamics of light-use efficiency in sun and shade leaves of Fagus sylvatica.

Sunflecks are transient patches of direct radiation that provide a substantial proportion of the daily irradiance to leaves in the lower canopy. In this position, faster photosynthetic induction would allow for higher sunfleck-use efficiency, as is commonly reported in the literature. Yet, when sunflecks are too few and far between, it may be more beneficial for shade leaves to prioritize efficient photosynthesis under shade. We investigated the temporal dynamics of photosynthetic induction, recovery under shade, and stomatal movement during a sunfleck, in sun and shade leaves of Fagus sylvatica from three provenances of contrasting origin. We found that shade leaves complete full induction in a shorter time than sun leaves, but that sun leaves respond faster than shade leaves due to their much larger amplitude of induction. The core-range provenance achieved faster stomatal opening in shade leaves, which may allow for better sunfleck-use efficiency in denser canopies and lower canopy positions. Our findings represent a paradigm shift for future research into light fluctuations in canopies, drawing attention to the ubiquitous importance of sunflecks for photosynthesis, not only in lower-canopy leaves where shade is prevalent, but particularly in the upper canopy where longer sunflecks are more common due to canopy openness.

Out of Africa: characterizing the natural variation in dynamic photosynthetic traits in a diverse population of African rice (Oryza glaberrima).

African rice (Oryza glaberrima) has adapted to challenging environments and is a promising source of genetic variation. We analysed dynamics of photosynthesis and morphology in a reference set of 155 O. glaberrima accessions. Plants were grown in an agronomy glasshouse to late tillering stage. Photosynthesis induction from darkness and the decrease in low light was measured by gas exchange and chlorophyll fluorescence along with root and shoot biomass, stomatal density, and leaf area. Steady-state and kinetic responses were modelled. We describe extensive natural variation in O. glaberrima for steady-state, induction, and reduction responses of photosynthesis that has value for gene discovery and crop improvement. Principal component analyses indicated key clusters of plant biomass, kinetics of photosynthesis (CO2 assimilation, A), and photoprotection induction and reduction (measured by non-photochemical quenching, NPQ), consistent with diverse adaptation. Accessions also clustered according to countries with differing water availability, stomatal conductance (gs), A, and NPQ, indicating that dynamic photosynthesis has adaptive value in O. glaberrima. Kinetics of NPQ, A, and gs showed high correlation with biomass and leaf area. We conclude that dynamic photosynthetic traits and NPQ are important within O. glaberrima, and we highlight NPQ kinetics and NPQ under low light.

Dynamic non-photochemical quenching in plants: from molecular mechanism to productivity.

Photoprotection refers to a set of well defined plant processes that help to prevent the deleterious effects of high and excess light on plant cells, especially within the chloroplast. Molecular components of chloroplast photoprotection are closely aligned with those of photosynthesis and together they influence productivity. Proof of principle now exists that major photoprotective processes such as non-photochemical quenching (NPQ) directly determine whole canopy photosynthesis, biomass and yield via prevention of photoinhibition and a momentary downregulation of photosynthetic quantum yield. However, this phenomenon has neither been quantified nor well characterized across different environments. Here we address this problem by assessing the existing literature with a different approach to that taken previously, beginning with our understanding of the molecular mechanism of NPQ and its regulation within dynamic environments. We then move to the leaf and the plant level, building an understanding of the circumstances (when and where) NPQ limits photosynthesis and linking to our understanding of how this might take place on a molecular and metabolic level. We argue that such approaches are needed to fine tune the relevant features necessary for improving dynamic NPQ in important crop species.

Rapid temperature responses of photosystem II efficiency forecast genotypic variation in rice vegetative heat tolerance.

A key target for the improvement of Oryza sativa (rice) is the development of heat-tolerant varieties. This necessitates the development of high-throughput methodologies for the screening of heat tolerance. Progress has been made to this end via visual scoring and chlorophyll fluorescence; however, these approaches demand large infrastructural investments to expose large populations of adult plants to heat stress. To address this bottleneck, we investigated the response of the maximum quantum efficiency of photosystem II (PSII) to rapidly increasing temperatures in excised leaf segments of juvenile rice plants. Segmented models explained the majority of the observed variation in response. Coefficients from these models, i.e. critical temperature (Tcrit ) and the initial response (m1 ), were evaluated for their usability for forecasting adult heat tolerance, measured as the vegetative heat tolerance of adult rice plants through visual (stay-green) and chlorophyll fluorescence (ɸPSII) approaches. We detected substantial variation in heat tolerance of a randomly selected set of indica rice varieties. Both Tcrit and m1 were associated with measured heat tolerance in adult plants, highlighting their usability as high-throughput proxies. Variation in heat tolerance was associated with daytime respiration but not with photosynthetic capacity, highlighting a role for the non-photorespiratory release of CO2 in heat tolerance. To date, this represents the first published instance of genetic variation in these key gas-exchange traits being quantified in response to heat stress in a diverse set of rice accessions. These results outline an efficient strategy for screening heat tolerance and accentuate the need to focus on reduced rates of respiration to improve heat tolerance in rice.

Nocturnal stomatal conductance in wheat is growth-stage specific and shows genotypic variation.

Nocturnal stomatal conductance (gsn ) represents a significant source of water loss, with implications for metabolism, thermal regulation and water-use efficiency. With increasing nocturnal temperatures due to climate change, it is vital to identify and understand variation in the magnitude and responses of gsn in major crops. We assessed interspecific variation in gsn and daytime stomatal conductance (gs ) in a wild relative and modern spring wheat genotype. To investigate intraspecific variation, we grew six modern wheat genotypes and two landraces under well watered, simulated field conditions. For the diurnal data, higher gsn in the wild relative was associated with significantly lower nocturnal respiration and higher daytime CO2 assimilation while both species exhibited declines in gsn post-dusk and pre-dawn. Lifetime gsn achieved rates of 5.7-18.9% of gs . Magnitude of gsn was genotype specific 'and positively correlated with gs . gsn and gs were significantly higher on the adaxial surface. No relationship was determined between harvest characteristics, stomatal morphology and gsn , while cuticular conductance was genotype specific. Finally, for the majority of genotypes, gsn declined with age. Here we present the discovery that variation in gsn occurs across developmental, morphological and temporal scales in nonstressed wheat, presenting opportunities for exploiting intrinsic variation under heat or water stressed conditions.

The potential of resilient carbon dynamics for stabilizing crop reproductive development and productivity during heat stress.

Impaired carbon metabolism and reproductive development constrain crop productivity during heat stress. Reproductive development is energy intensive, and its requirement for respiratory substrates rises as associated metabolism increases with temperature. Understanding how these processes are integrated and the extent to which they contribute to the maintenance of yield during and following periods of elevated temperatures is important for developing climate-resilient crops. Recent studies are beginning to demonstrate links between processes underlying carbon dynamics and reproduction during heat stress, consequently a summation of research that has been reported thus far and an evaluation of purported associations are needed to guide and stimulate future research. To this end, we review recent studies relating to source-sink dynamics, non-foliar photosynthesis and net carbon gain as pivotal in understanding how to improve reproductive development and crop productivity during heat stress. Rapid and precise phenotyping during narrow phenological windows will be important for understanding mechanisms underlying these processes, thus we discuss the development of relevant high-throughput phenotyping approaches that will allow for more informed decision-making regarding future crop improvement.

The effect of canopy architecture on the patterning of "windflecks" within a wheat canopy.

Under field conditions, plants are subject to wind-induced movement which creates fluctuations of light intensity and spectral quality reaching the leaves, defined here as windflecks. Within this study, irradiance within two contrasting wheat (Triticum aestivum) canopies during full sun conditions was measured using a spectroradiometer to determine the frequency, duration and magnitude of low- to high-light events plus the spectral composition during wind-induced movement. Similarly, a static canopy was modelled using three-dimensional reconstruction and ray tracing to determine fleck characteristics without the presence of wind. Corresponding architectural traits were measured manually and in silico including plant height, leaf area and angle plus biomechanical properties. Light intensity can differ up to 40% during a windfleck, with changes occurring on a sub-second scale compared to ~5 min in canopies not subject to wind. Features such as a shorter height, more erect leaf stature and having an open structure led to an increased frequency and reduced time interval of light flecks in the CMH79A canopy compared to Paragon. This finding illustrates the potential for architectural traits to be selected to improve the canopy light environment and provides the foundation to further explore the links between plant form and function in crop canopies.

Field-based remote sensing models predict radiation use efficiency in wheat.

Wheat yields are stagnating or declining in many regions, requiring efforts to improve the light conversion efficiency, known as radiation use efficiency (RUE). RUE is a key trait in plant physiology because it links light capture and primary metabolism with biomass accumulation and yield, but its measurement is time consuming and this has limited its use in fundamental research and large-scale physiological breeding. In this study, high-throughput plant phenotyping (HTPP) approaches were used among a population of field-grown wheat with variation in RUE and photosynthetic traits to build predictive models of RUE, biomass, and intercepted photosynthetically active radiation (IPAR). Three approaches were used: best combination of sensors; canopy vegetation indices; and partial least squares regression. The use of remote sensing models predicted RUE with up to 70% accuracy compared with ground truth data. Water indices and canopy greenness indices [normalized difference vegetation index (NDVI), enhanced vegetation index (EVI)] are the better option to predict RUE, biomass, and IPAR, and indices related to gas exchange, non-photochemical quenching [photochemical reflectance index (PRI)] and senescence [structural-insensitive pigment index (SIPI)] are better predictors for these traits at the vegetative and grain-filling stages, respectively. These models will be instrumental to explain canopy processes, improve crop growth and yield modelling, and potentially be used to predict RUE in different crops or ecosystems.

Variation in key leaf photosynthetic traits across wheat wild relatives is accession dependent not species dependent.

The wild relatives of modern wheat represent an underutilized source of genetic and phenotypic diversity and are of interest in breeding owing to their wide adaptation to diverse environments. Leaf photosynthetic traits underpin the rate of production of biomass and yield and have not been systematically explored in the wheat relatives. This paper identifies and quantifies the phenotypic variation in photosynthetic, stomatal, and morphological traits in up to 88 wheat wild relative accessions across five genera. Both steady-state measurements and dynamic responses to step changes in light intensity are assessed. A 2.3-fold variation for flag leaf light and CO2 -saturated rates of photosynthesis Amax was observed. Many accessions showing higher and more variable Amax , maximum rates of carboxylation, electron transport, and Rubisco activity when compared with modern genotypes. Variation in dynamic traits was also significant; with distinct genus-specific trends in rates of induction of nonphotochemical quenching and rate of stomatal opening. We conclude that utilization of wild relatives for improvement of photosynthesis is supported by the existence of a high degree of natural variation in key traits and should consider not only genus-level properties but variation between individual accessions.

Recovering Wind-Induced Plant Motion in Dense Field Environments via Deep Learning and Multiple Object Tracking.

Understanding the relationships between local environmental conditions and plant structure and function is critical for both fundamental science and for improving the performance of crops in field settings. Wind-induced plant motion is important in most agricultural systems, yet the complexity of the field environment means that it remained understudied. Despite the ready availability of image sequences showing plant motion, the cultivation of crop plants in dense field stands makes it difficult to detect features and characterize their general movement traits. Here, we present a robust method for characterizing motion in field-grown wheat plants (Triticum aestivum) from time-ordered sequences of red, green, and blue images. A series of crops and augmentations was applied to a dataset of 290 collected and annotated images of ear tips to increase variation and resolution when training a convolutional neural network. This approach enables wheat ears to be detected in the field without the need for camera calibration or a fixed imaging position. Videos of wheat plants moving in the wind were also collected and split into their component frames. Ear tips were detected using the trained network, then tracked between frames using a probabilistic tracking algorithm to approximate movement. These data can be used to characterize key movement traits, such as periodicity, and obtain more detailed static plant properties to assess plant structure and function in the field. Automated data extraction may be possible for informing lodging models, breeding programs, and linking movement properties to canopy light distributions and dynamic light fluctuation.

Photoprotective energy dissipation is greater in the lower, not the upper, regions of a rice canopy: a 3D analysis.

High light intensities raise photosynthetic and plant growth rates but can cause damage to the photosynthetic machinery. The likelihood and severity of deleterious effects are minimised by a set of photoprotective mechanisms, one key process being the controlled dissipation of energy from chlorophyll within PSII known as non-photochemical quenching (NPQ). Although ubiquitous, the role of NPQ in plant productivity is important because it momentarily reduces the quantum efficiency of photosynthesis. Rice plants overexpressing and deficient in the gene encoding a central regulator of NPQ, the protein PsbS, were used to assess the effect of protective effectiveness of NPQ (pNPQ) at the canopy scale. Using a combination of three-dimensional reconstruction, modelling, chlorophyll fluorescence, and gas exchange, the influence of altered NPQ capacity on the distribution of pNPQ was explored. A higher phototolerance in the lower layers of a canopy was found, regardless of genotype, suggesting a mechanism for increased protection for leaves that experience relatively low light intensities interspersed with brief periods of high light. Relative to wild-type plants, psbS overexpressors have a reduced risk of photoinactivation and early growth advantage, demonstrating that manipulating photoprotective mechanisms can impact both subcellular mechanisms and whole-canopy function.

Rice with reduced stomatal density conserves water and has improved drought tolerance under future climate conditions.

Much of humanity relies on rice (Oryza sativa) as a food source, but cultivation is water intensive and the crop is vulnerable to drought and high temperatures. Under climate change, periods of reduced water availability and high temperature are expected to become more frequent, leading to detrimental effects on rice yields. We engineered the high-yielding rice cultivar 'IR64' to produce fewer stomata by manipulating the level of a developmental signal. We overexpressed the rice epidermal patterning factor OsEPF1, creating plants with substantially reduced stomatal density and correspondingly low stomatal conductance. Low stomatal density rice lines were more able to conserve water, using c. 60% of the normal amount between weeks 4 and 5 post germination. When grown at elevated atmospheric CO2 , rice plants with low stomatal density were able to maintain their stomatal conductance and survive drought and high temperature (40°C) for longer than control plants. Low stomatal density rice gave equivalent or even improved yields, despite a reduced rate of photosynthesis in some conditions. Rice plants with fewer stomata are drought tolerant and more conservative in their water use, and they should perform better in the future when climate change is expected to threaten food security.

Suboptimal Acclimation of Photosynthesis to Light in Wheat Canopies.

Photosynthetic acclimation (photoacclimation) is the process whereby leaves alter their morphology and/or biochemistry to optimize photosynthetic efficiency and productivity according to long-term changes in the light environment. The three-dimensional architecture of plant canopies imposes complex light dynamics, but the drivers for photoacclimation in such fluctuating environments are poorly understood. A technique for high-resolution three-dimensional reconstruction was combined with ray tracing to simulate a daily time course of radiation profiles for architecturally contrasting field-grown wheat (Triticum aestivum) canopies. An empirical model of photoacclimation was adapted to predict the optimal distribution of photosynthesis according to the fluctuating light patterns throughout the canopies. While the photoacclimation model output showed good correlation with field-measured gas-exchange data at the top of the canopy, it predicted a lower optimal light-saturated rate of photosynthesis at the base. Leaf Rubisco and protein contents were consistent with the measured optimal light-saturated rate of photosynthesis. We conclude that, although the photosynthetic capacity of leaves is high enough to exploit brief periods of high light within the canopy (particularly toward the base), the frequency and duration of such sunflecks are too small to make acclimation a viable strategy in terms of carbon gain. This suboptimal acclimation renders a large portion of residual photosynthetic capacity unused and reduces photosynthetic nitrogen use efficiency at the canopy level, with further implications for photosynthetic productivity. It is argued that (1) this represents an untapped source of photosynthetic potential and (2) canopy nitrogen could be lowered with no detriment to carbon gain or grain protein content.