Pollinator-assisted plant phenotyping, selection, and breeding for crop resilience to abiotic stresses.

Food security is threatened by climate change, with heat and drought being the main stresses affecting crop physiology and ecosystem services, such as plant-pollinator interactions. We hypothesize that tracking and ranking pollinators' preferences for flowers under environmental pressure could be used as a marker of plant quality for agricultural breeding to increase crop stress tolerance. Despite increasing relevance of flowers as the most stress sensitive organs, phenotyping platforms aim at identifying traits of resilience by assessing the plant physiological status through remote sensing-assisted vegetative indexes, but find strong bottlenecks in quantifying flower traits and in accurate genotype-to-phenotype prediction. However, as the transport of photoassimilates from leaves (sources) to flowers (sinks) is reduced in low-resilient plants, flowers are better indicators than leaves of plant well-being. Indeed, the chemical composition and amount of pollen and nectar that flowers produce, which ultimately serve as food resources for pollinators, change in response to environmental cues. Therefore, pollinators' preferences could be used as a measure of functional source-to-sink relationships for breeding decisions. To achieve this challenging goal, we propose to develop a pollinator-assisted phenotyping and selection platform for automated quantification of Genotype × Environment × Pollinator interactions through an insect geo-positioning system. Pollinator-assisted selection can be validated by metabolic, transcriptomic, and ionomic traits, and mapping of candidate genes, linking floral and leaf traits, pollinator preferences, plant resilience, and crop productivity. This radical new approach can change the current paradigm of plant phenotyping and find new paths for crop redomestication and breeding assisted by ecological decisions.

Auxin regulates source-sink carbohydrate partitioning and reproductive organ development in rice.

Carbohydrate partitioning between the source and sink tissues plays an important role in regulating plant growth and development. However, the molecular mechanisms regulating this process remain poorly understood. In this study, we show that elevated auxin levels in the rice dao mutant cause increased accumulation of sucrose in the photosynthetic leaves but reduced sucrose content in the reproductive organs (particularly in the lodicules, anthers, and ovaries), leading to closed spikelets, indehiscent anthers, and parthenocarpic seeds. RNA sequencing analysis revealed that the expression of AUXIN RESPONSE FACTOR 18 (OsARF18) and OsARF2 is significantly up- and down-regulated, respectively, in the lodicule of dao mutant. Overexpression of OsARF18 or knocking out of OsARF2 phenocopies the dao mutant. We demonstrate that OsARF2 regulates the expression of OsSUT1 through direct binding to the sugar-responsive elements (SuREs) in the OsSUT1 promoter and that OsARF18 represses the expression of OsARF2 and OsSUT1 via direct binding to the auxin-responsive element (AuxRE) or SuRE in their promoters, respectively. Furthermore, overexpression of OsSUT1 in the dao and Osarf2 mutant backgrounds could largely rescue the spikelets' opening and seed-setting defects. Collectively, our results reveal an auxin signaling cascade regulating source-sink carbohydrate partitioning and reproductive organ development in rice.

Efficient sugar utilization and transition from oxidative to substrate-level phosphorylation in high starch storage roots of African cassava genotypes.

Cassava's storage roots represent one of the most important sources of nutritional carbohydrates worldwide. Particularly, smallholder farmers in sub-Saharan Africa depend on this crop plant, where resilient and yield-improved varieties are of vital importance to support steadily increasing populations. Aided by a growing understanding of the plant's metabolism and physiology, targeted improvement concepts already led to visible gains in recent years. To expand our knowledge and to contribute to these successes, we investigated storage roots of eight cassava genotypes with differential dry matter content from three successive field trials for their proteomic and metabolic profiles. At large, the metabolic focus in storage roots transitioned from cellular growth processes toward carbohydrate and nitrogen storage with increasing dry matter content. This is reflected in higher abundance of proteins related to nucleotide synthesis, protein turnover, and vacuolar energization in low starch genotypes, while proteins involved in sugar conversion and glycolysis were more prevalent in high dry matter genotypes. This shift in metabolic orientation was underlined by a clear transition from oxidative- to substrate-level phosphorylation in high dry matter genotypes. Our analyses highlight metabolic patterns that are consistently and quantitatively associated with high dry matter accumulation in cassava storage roots, providing fundamental understanding of cassava's metabolism as well as a data resource for targeted genetic improvement.

Root-to-shoot signaling positively mediates source-sink relation in late growth stages in diploid and tetraploid wheat.

Non-hydraulic root source signaling (nHRS) is a unique positive response to soil drying in the regulation of plant growth and development. However, it is unclear how the nHRS mediates the tradeoff between source and sink at the late growth stages and its adaptive mechanisms in primitive wheat. To address this issue, a root-splitting design was made by inserting solid partition in the middle of the pot culture to induce the occurrence of nHRS using four wheat cultivars (MO1 and MO4, diploid; DM22 and DM31, tetraploid) as materials. Three water treatments were designed as 1) both halves watered (CK), 2) holistic root system watered then droughted (FS), 3) one-half of the root system watered and half droughted (PS). FS and PS were designed to compare the role of the full root system and split root system to induce nHRS. Leaves samples were collected during booting and anthesis to compare the role of nHRS at both growth stages. The data indicated that under PS treatment, ABA concentration was significantly higher than FS and CK, demonstrating the induction of nHRS in split root design and nHRS decreased cytokinin (ZR) levels, particularly in the PS treatment. Soluble sugar and proline accumulation were higher in the anthesis stage as compared to the booting stage. POD activity was higher at anthesis, while CAT was higher at the booting stage. Increased ABA (nHRS) correlated with source-sink relationships and metabolic rate (i.e., leaf) connecting other stress signals. Biomass density showed superior resource acquisition and utilization capabilities in both FS and PS treatment as compared to CK in all plants. Our findings indicate that nHRS-induced alterations in phytohormones and their effect on source-sink relations were allied with the growth stages in primitive wheat.

Tissue- and time-dependent metabolite profiles during early grain development under normal and high night-time temperature conditions.

BACKGROUND: Wheat grain development in the first few days after pollination determines the number of endosperm cells that influence grain yield potential and is susceptible to various environmental conditions, including high night temperatures (HNTs). Flag leaves and seed-associated bracts (glumes, awn, palea, and lemma) provide nutrients to the developing seed. However, the specific metabolic roles of these tissues are uncertain, especially their dynamics at different developmental stages and the time in a day. Tissue- and time-dependent metabolite profiling may hint at the metabolic roles of tissues and the mechanisms of how HNTs affect daytime metabolic status in early grain development. RESULTS: The metabolite profiles of flag leaf, bract, seed (embryo and endosperm), and entire spike were analyzed at 12:00 (day) and 23:00 (night) on 2, 4, and 6 days after fertilization under control and HNT conditions. The metabolite levels in flag leaves and bracts showed day/night oscillations, while their behaviors were distinct between the tissues. Some metabolites, such as sucrose, cellobiose, and succinic acid, showed contrasting oscillations in the two photosynthetic tissues. In contrast, seed metabolite levels differed due to the days after fertilization rather than the time in a day. The seed metabolite profile altered earlier in the HNT than in the control condition, likely associated with accelerated grain development caused by HNT. HNT also disrupted the day/night oscillation of sugar accumulation in flag leaves and bracts. CONCLUSIONS: These results highlight distinct metabolic roles of flag leaves and bracts during wheat early seed development. The seed metabolite levels are related to the developmental stages. The early metabolic events in the seeds and the disruption of the day/night metabolic cycle in photosynthetic tissues may partly explain the adverse effects of HNT on grain yield.

Grain yield trade-offs in spike-branching wheat can be mitigated by elite alleles affecting sink capacity and post-anthesis source activity.

Introducing variations in inflorescence architecture, such as the 'Miracle-Wheat' (Triticum turgidum convar. compositum (L.f.) Filat.) with a branching spike, has relevance for enhancing wheat grain yield. However, in the spike-branching genotypes, the increase in spikelet number is generally not translated into grain yield advantage because of reduced grains per spikelet and grain weight. Here, we investigated if such trade-offs might be a function of source-sink strength by using 385 recombinant inbred lines developed by intercrossing the spike-branching landrace TRI 984 and CIRNO C2008, an elite durum (T. durum L.) cultivar; they were genotyped using the 25K array. Various plant and spike architectural traits, including flag leaf, peduncle, and spike senescence rate, were phenotyped under field conditions for 2 consecutive years. On chromosome 5AL, we found a new modifier QTL for spike branching, branched headt3 (bht-A3), which was epistatic to the previously known bht-A1 locus. Besides, bht-A3 was associated with more grains per spikelet and a delay in flag leaf senescence rate. Importantly, favourable alleles, viz. bht-A3 and grain protein content (gpc-B1) that delayed senescence, are required to improve grain number and grain weight in the spike-branching genotypes. In summary, achieving a balanced source-sink relationship might minimize grain yield trade-offs in Miracle-Wheat.

Non-foliar photosynthesis and nitrogen assimilation influence grain yield in durum wheat regardless of water conditions.

There is a need to generate improved crop varieties adapted to the ongoing changes in the climate. We studied durum wheat canopy and central metabolism of six different photosynthetic organs in two yield-contrasting varieties. The aim was to understand the mechanisms associated with the water stress response and yield performance. Water stress strongly reduced grain yield, plant biomass, and leaf photosynthesis, and down-regulated C/N-metabolism genes and key protein levels, which occurred mainly in leaf blades. By contrast, higher yield was associated with high ear dry weight and lower biomass and ears per area, highlighting the advantage of reduced tillering and the consequent improvement in sink strength, which promoted C/N metabolism at the whole plant level. An improved C metabolism in blades and ear bracts and N assimilation in all photosynthetic organs facilitated C/N remobilization to the grain and promoted yield. Therefore, we propose that further yield gains in Mediterranean conditions could be achieved by considering the source-sink dynamics and the contribution of non-foliar organs, and particularly N assimilation and remobilization during the late growth stages. We highlight the power of linking phenotyping with plant metabolism to identify novel traits at the whole plant level to support breeding programmes.

Spatial heterogeneity of biomass turnover has contrasting effects on synchrony and stability in trophic metacommunities.

Spatial heterogeneity is a fundamental feature of ecosystems, and ecologists have identified it as a factor promoting the stability of population dynamics. In particular, differences in interaction strengths and resource supply between patches generate an asymmetry of biomass turnover with a fast and a slow patch coupled by a mobile predator. Here, we demonstrate that asymmetry leads to opposite stability patterns in metacommunities receiving localized perturbations depending on the characteristics of the perturbed patch. Perturbing prey in the fast patch synchronizes the dynamics of prey biomass between the two patches and destabilizes predator dynamics by increasing the predator's temporal variability. Conversely, perturbing prey in the slow patch decreases the synchrony of the prey's dynamics and stabilizes predator dynamics. Our results have implications for conservation ecology and suggest reinforcing protection policies in fast patches to dampen the effects of perturbations and promote the stability of population dynamics at the regional scale.

The ecological causes of functional distinctiveness in communities.

Recent work has shown that evaluating functional trait distinctiveness, the average trait distance of a species to other species in a community offers promising insights into biodiversity dynamics and ecosystem functioning. However, the ecological mechanisms underlying the emergence and persistence of functionally distinct species are poorly understood. Here, we address the issue by considering a heterogeneous fitness landscape whereby functional dimensions encompass peaks representing trait combinations yielding positive population growth rates in a community. We identify four ecological cases contributing to the emergence and persistence of functionally distinct species. First, environmental heterogeneity or alternative phenotypic designs can drive positive population growth of functionally distinct species. Second, sink populations with negative population growth can deviate from local fitness peaks and be functionally distinct. Third, species found at the margin of the fitness landscape can persist but be functionally distinct. Fourth, biotic interactions (positive or negative) can dynamically alter the fitness landscape. We offer examples of these four cases and guidelines to distinguish between them. In addition to these deterministic processes, we explore how stochastic dispersal limitation can yield functional distinctiveness. Our framework offers a novel perspective on the relationship between fitness landscape heterogeneity and the functional composition of ecological assemblages.

The Alteration of Tomato Chloroplast Vesiculation Positively Affects Whole-Plant Source-Sink Relations and Fruit Metabolism under Stress Conditions.

Changes in climate conditions can negatively affect the productivity of crop plants. They can induce chloroplast degradation (senescence), which leads to decreased source capacity, as well as decreased whole-plant carbon/nitrogen assimilation and allocation. The importance, contribution and mechanisms of action regulating source-tissue capacity under stress conditions in tomato (Solanum lycopersicum) are not well understood. We hypothesized that delaying chloroplast degradation by altering the activity of the tomato chloroplast vesiculation (CV) under stress would lead to more efficient use of carbon and nitrogen and to higher yields. Tomato CV is upregulated under stress conditions. Specific induction of CV in leaves at the fruit development stage resulted in stress-induced senescence and negatively affected fruit yield, without any positive effects on fruit quality. Clustered Regularly Interspaced Short Palindromic Repeats/CRISPR-associated protein 9 (CRISPR/CAS9) knockout CV plants, generated using a near-isogenic tomato line with enhanced sink capacity, exhibited stress tolerance at both the vegetative and the reproductive stages, leading to enhanced fruit quantity, quality and harvest index. Detailed metabolic and transcriptomic network analysis of sink tissue revealed that the l-glutamine and l-arginine biosynthesis pathways are associated with stress-response conditions and also identified putative novel genes involved in tomato fruit quality under stress. Our results are the first to demonstrate the feasibility of delayed stress-induced senescence as a stress-tolerance trait in a fleshy fruit crop, to highlight the involvement of the CV pathway in the regulation of source strength under stress and to identify genes and metabolic pathways involved in increased tomato sink capacity under stress conditions.

Analysis of durum wheat photosynthetic organs during grain filling reveals the ear as a water stress-tolerant organ and the peduncle as the largest pool of primary metabolites.

MAIN CONCLUSION: The pool of carbon- and nitrogen-rich metabolites is quantitatively relevant in non-foliar photosynthetic organs during grain filling, which have a better response to water limitation than flag leaves. The response of durum wheat to contrasting water regimes has been extensively studied at leaf and agronomic level in previous studies, but the water stress effects on source-sink dynamics, particularly non-foliar photosynthetic organs, is more limited. Our study aims to investigate the response of different photosynthetic organs to water stress and to quantify the pool of carbon and nitrogen metabolites available for grain filling. Five durum wheat varieties were grown in field trials in the Spanish region of Castile and León under irrigated and rainfed conditions. Water stress led to a significant decrease in yield, biomass, and carbon and nitrogen assimilation, improved water use efficiency, and modified grain quality traits in the five varieties. The pool of carbon (glucose, glucose-6-phosphate, fructose, sucrose, starch, and malate) and nitrogen (glutamate, amino acids, proteins and chlorophylls) metabolites in leaf blades and sheaths, peduncles, awns, glumes and lemmas were also analysed. The results showed that the metabolism of the blades and peduncles was the most susceptible to water stress, while ear metabolism showed higher stability, particularly at mid-grain filling. Interestingly, the total metabolite content per organ highlighted that a large source of nutrients, which may be directly involved in grain filling, are found outside the blades, with the peduncles being quantitatively the most relevant. We conclude that yield improvements in our Mediterranean agro-ecosystem are highly linked to the success of shoots in producing ears and a higher number of grains, while grain filling is highly dependent on the capacity of non-foliar organs to fix CO2 and N. The ear organs show higher stress resilience than other organs, which deserves our attention in future breeding programmes.

Physiological implications of SWEETs in plants and their potential applications in improving source-sink relationships for enhanced yield.

The sugars will eventually be exported transporters (SWEET) family of transporters in plants is identified as a novel class of sugar carriers capable of transporting sugars, sugar alcohols and hormones. Functioning in intercellular sugar transport, SWEETs influence a wide range of physiologically important processes. SWEETs regulate the development of sink organs by providing nutritional support from source leaves, responses to abiotic stresses by maintaining intracellular sugar concentrations, and host-pathogen interactions through the modulation of apoplastic sugar levels. Many bacterial and fungal pathogens activate the expression of SWEET genes in species such as rice and Arabidopsis to gain access to the nutrients that support virulence. The genetic manipulation of SWEETs has led to the generation of bacterial blight (BB)-resistant rice varieties. Similarly, while the overexpression of the SWEETs involved in sucrose export from leaves and pathogenesis led to growth retardation and yield penalties, plants overexpressing SWEETs show improved disease resistance. Such findings demonstrate the complex functions of SWEETs in growth and stress tolerance. Here, we review the importance of SWEETs in plant-pathogen and source-sink interactions and abiotic stress resistance. We highlight the possible applications of SWEETs in crop improvement programmes aimed at improving sink and source strengths important for enhancing the sustainability of yield. We discuss how the adverse effects of the overexpression of SWEETs on plant growth may be overcome.

Modified source-sink dynamics govern resource exchange in ectomycorrhizal symbiosis.

Ectomycorrhizal symbiosis between roots and fungi is founded on the movement of carbon from plants to fungi, and of soil resources from fungi to plants. Framing this movement as a trade can facilitate an understanding of how this mutualism has developed over evolutionary time, but fails to explain experimental observations of carbon and nutrient movement. Here, I propose that source-sink dynamics are an essential basic model to explain the movement of plant and fungal resources, which may be modified by plant immune response, variability in fungal molecular repertoires, and competition in the soil. Source-sink dynamics provide testable hypotheses to illuminate mechanisms of ectomycorrhizal resource movement and its consequences for mutualism stability and forest function under climate change.

Do stomata optimize turgor-driven growth? A new framework for integrating stomata response with whole-plant hydraulics and carbon balance.

Every existing optimal stomatal model uses photosynthetic carbon assimilation as a proxy for plant evolutionary fitness. However, assimilation and growth are often decoupled, making assimilation less ideal for representing fitness when optimizing stomatal conductance to water vapor and carbon dioxide. Instead, growth should be considered a closer proxy for fitness. We hypothesize stomata have evolved to maximize turgor-driven growth, instead of assimilation, over entire plants' lifetimes, improving their abilities to compete and reproduce. We develop a stomata model that dynamically maximizes whole-stem growth following principles from turgor-driven growth models. Stomata open to assimilate carbohydrates that supply growth and osmotically generate turgor, while stomata close to prevent losses of turgor and growth due to negative water potentials. In steady state, the growth optimization model captures realistic stomatal, growth, and carbohydrate responses to environmental cues, reconciles conflicting interpretations within existing stomatal optimization theories, and explains patterns of carbohydrate storage and xylem conductance observed during and after drought. Our growth optimization hypothesis introduces a new paradigm for stomatal optimization models, elevates the role of whole-plant carbon use and carbon storage in stomatal functioning, and has the potential to simultaneously predict gross productivity, net productivity, and plant mortality through a single, consistent modeling framework.

Low light stress promotes new tiller regeneration by changing source-sink relationship and activating expression of expansin genes in wheat.

Low light stress seriously decreased wheat grain number through the formation of aborted spike during the reproductive period and induced new tiller regeneration to offset the loss of grain number. However, the mechanism by which plants coordinate spike aborted growth and the regeneration of new tillers remains unknown. To better understand this coordinated process, morphological, physiological and transcriptomic analyses were performed under low light stress at the young microspore stage. Our findings indicated that leaves exhausted most stored carbohydrates in 1 day of darkness. However, spike and uppermost internode (UI) were converted from sink to source, due to increased abscisic acid (ABA) content and decreased cytokinin content. During this process, genes encoding amylases, Sugars Will Eventually be Exported Transporters (SWEET) and sucrose transporters or sucrose carriers (SUT/SUC) were upregulated in spike and UI, which degraded starch into soluble sugars and loaded them into the phloem. Subsequently, soluble sugars were transported to tiller node (TN) where cytokinin and auxin content increased and ABA content decreased, followed by unloading into TN cells by upregulated cell wall invertase (CWINV) genes and highly expressed H+ /hexose symporter genes. Finally, expansin genes integrated the sugar pathway and hormone pathway, and regulate the formation of new tillers directly.

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.

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.

Bulb growth potential is independent of leaf longevity for the spring ephemeral Erythronium americanum Ker-Gawl.

Growth in most spring ephemerals is decreased under warmer temperatures. Although photosynthetic activities are improved at warmer temperatures, leaves senesce earlier, which prevents the bulb from reaching a larger size. A longer leaf life duration during a warm spring, therefore, may improve bulb mass. We tested this hypothesis by modulating leaf life span of Erythronium americanum through the application of Promalin® (PRO; cytokinins and gibberellins) that prolonged or silver thiosulfate (STS) that reduced leaf duration. Gas exchange and chlorophyll fluorescence were measured along with leaf and bulb carbohydrate concentrations. Plants were also pulse labelled with 13CO2 to monitor sugar transport to the bulb. Lower photosynthetic rates and shorter leaf life span of STS plants reduced the amount of carbon that they assimilated during the season, resulting in a smaller bulb compared with control plants. PRO plants maintained their photosynthetic rates for a longer period than control plants, yet final bulb biomass did not differ between them. We conclude that seasonal growth for E. americanum is not limited by leaf life duration under warm growing conditions, but rather by limited sink growth capacity. Under global warming, spring geophytes might be at risk of being reduced in size and, eventually, reproducing less frequently.

Crops under past diversification and ongoing climate change: more than just selection of nuclear genes for flowering.

Diversification and breeding following domestication and under current climate change across the globe are the two most significant evolutionary events experienced by major crops. Diversification of crops from their wild ancestors has favored dramatic changes in the sensitivity of the plants to the environment, particularly significantly in transducing light inputs to the circadian clock, which has allowed the growth of major crops in the relatively short growing season experienced in the Northern Hemisphere. Historically, mutants and the mapping of quantitative trait loci (QTL) have facilitated the identification and the cloning of genes that underlie major changes of the clock and the regulation of flowering. Recent studies have suggested that the thermal plasticity of the circadian clock output, and not just the core genes that follow temperature compensation, has also been under selection during diversification and breeding. Wild alleles that accelerate output rhythmicity could be beneficial for crop resilience. Furthermore, wild alleles with beneficial and flowering-independent effects under stress indicate their possible role in maintaining a balanced source-sink relationship, thereby allowing productivity under climatic change. Because the chloroplast genome also regulates the plasticity of the clock output, mapping populations including cytonuclear interactions should be utilized within an integrated field and clock phenomics framework. In this review, we highlight the need to integrate physiological and developmental approaches (physio-devo) to gain a better understanding when re-domesticating wild gene alleles into modern cultivars to increase their robustness under abiotic heat and drought stresses.

Negative relationship between photosynthesis and late-stage canopy development and senescence over Tibetan Plateau.

Canopy greening, which is associated with significant canopy structure changes, is the most notable signal of ecosystem changes in response to anthropogenic climate change. However, our knowledge of the changing pattern of canopy development and senescence, and its endogenous and climatic drivers is still limited. Here, we used the Normalized Difference Vegetation Index (NDVI) to quantify the changes in the speed of canopy development and senescence over the Tibetan Plateau (TP) during 2000-2018, and used a solar-induced chlorophyll fluorescence dataset as a proxy for photosynthesis, in combination with climate datasets to decipher the endogenous and climatic drivers of the interannual variation in canopy changes. We found that the canopy development during the early green-up stage (April-May) is accelerating at a rate of 0.45-0.8 × 10-3  month-1  year-1 . However, this accelerating canopy development was largely offset by a decelerating canopy development during June and July (-0.61 to -0.51 × 10-3  month-1  year-1 ), leading to the peak NDVI over the TP increasing at a rate of only one fifth of that in northern temperate regions, and less than one tenth of that in the Arctic and boreal regions. During the green-down period, we observed a significant accelerating canopy senescence during October. Photosynthesis was found to be the dominant driver for canopy changes over the TP. Increasing photosynthesis stimulates canopy development during the early green-up stage. However, slower canopy development and accelerated senescence was found with larger photosynthesis in late growth stages. This negative relationship between photosynthesis and canopy development is probably linked to the source-sink balance of plants and shifts in the allocation regime. These results suggest a sink limitation for plant growth over the TP. The impact of canopy greening on the carbon cycle may be more complicated than the source-oriented paradigm used in current ecosystem models.