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In vegetation stands, plants receive red to far‐red ratio (R:FR) signals of varying strength from all directions. However, plant responses to variations in R:FR reflected from below have been largely ignored despite their potential consequences for plant performance. Using a heterogeneous rose canopy, which consists of bent shoots down in the canopy and vertically growing upright shoots, we quantified upward far‐red reflection by bent shoots and its consequences for upright shoot architecture. With a three‐dimensional plant model, we assessed consequences of responses to R:FR from below for plant photosynthesis. Bent shoots reflected substantially more far‐red than red light, causing reduced R:FR in light reflected upwards. Leaf inclination angles increased in upright shoots which received low R:FR reflected from below. The increased leaf angle led to an increase in simulated plant photosynthesis only when this low R:FR was reflected off their own bent shoots and not when it reflected off neighbour bent shoots. We conclude that plant response to R:FR from below is an under‐explored phenomenon which may have contrasting consequences for plant performance depending on the type of vegetation or crop system. The responses are beneficial for performance only when R:FR is reflected by lower foliage of the same plants., In vegetation stands, plants receive light signals of varying strength from all directions. The presence of lower canopy decreased the red to far‐red ratio (R:FR) in light reflected upwards, which induced a steeper leaf angle in the upper canopy. Such a response to R:FR from below, when reflecting from the lower part of the same canopy, increased whole canopy photosynthesis by allowing more light to penetrate to the lower canopy.

Background and aims UVB radiation can rapidly induce gene regulation leading to cumulative changes for plant physiology and morphology. We hypothesized that a transgenerational effect of chronic exposure to solar short UV modulates the offspring’s responses to UVB and blue light, and that the transgenerational effect is genotype dependent. Methods We established a factorial experiment combining two Vicia faba L. accessions, two parental UV treatments (full sunlight and exclusion of short UV, 290−350 nm), and four offspring light treatments from the factorial combination of UVB and blue light. The accessions were Aurora from southern Sweden, and ILB938 from Andean region of Colombia and Ecuador. Key results The transgenerational effect influenced morphological responses to blue light differently in the two accessions. In Aurora, when UVB was absent, blue light increased shoot dry mass only in plants whose parents were protected from short UV. In ILB938, blue light increased leaf area and shoot dry mass more in plants whose parents were exposed to short UV than those that were not. Moreover, when the offspring was exposed to UVB, the transgenerational effect decreased in ILB938 and disappeared in Aurora. For flavonoids, the transgenerational effect was detected only in Aurora: parental exposure to short UV was associated with a greater induction of total quercetin in response to UVB. Transcript abundance was higher in Aurora than in ILB938 for both CHALCONE SYNTHASE (99-fold) and DON-GLUCOSYLTRANSFERASE 1 (19-fold). Conclusions The results supported both hypotheses. Solar short UV had transgenerational effects on progeny responses to blue and UVB radiation, and they differed between the accessions. These transgenerational effects could be adaptive by acclimation of slow and cumulative morphological change, and by early build-up of UV protection through flavonoid accumulation on UVB exposure. The differences between the two accessions aligned with their adaptation to contrasting UV environments.

Although plants are sessile organisms, they can modulate their phenotype so as to cope with environmental stresses such as herbivore attack and competition with neighbouring plants. Plant-produced volatile compounds mediate various aspects of plant defence. The emission of volatiles has costs and benefits. Research on the role of plant volatiles in defence has focused primarily on the responses of individual plants. However, in nature, plants rarely occur as isolated individuals but are members of plant communities where they compete for resources and exchange information with other plants. In this review, we address the effects of neighbouring plants on plant volatile-mediated defences. We will outline the various roles of volatile compounds in the interactions between plants and other organisms, address the mechanisms of plant neighbour perception in plant communities, and discuss how neighbour detection and volatile signalling are interconnected. Finally, we will outline the most urgent questions to be addressed in the future.

International audience; Oilseed rape (Brassica napus L.) is a crop with a complex aerial architecture that can cause self-shading leading to a vertical light gradient over the foliage. Mutual shading between neighboring plants at a high sowing density also results in an alteration of photosynthetically active radiation (PAR) absorption by lower leaves. The aim of this study was to analyze the impact that light restriction on lower leaves has on shoot architecture, biomass production and allocation, nitrogen (N) fluxes, and progression of sequential senescence. Field-grown plants were collected at the end of the vegetative rest period and grown in hydroponic conditions until pod maturity. A shading treatment corresponding to a 43.4 % reduction of PAR was applied at the early flowering stage. N uptake and fluxes of N allocation and remobilization were determined by supplying (KNO3)-N-15 in the nutrient solution. Photosynthesis and expression of SAG12 and Cab genes (indicators of leaf senescence progression) were also analyzed on different leaf ranks. The results showed that shading enhanced leaf development on the main stem and ramifications to optimize light capture. The expression pattern of the SAG12/Cab molecular indicator suggested a delay in leaf senescence that allowed leaf life span to be extended resulting in a more efficient leaf compound remobilization, with lower N residual contents in fallen leaves under shading. N uptake increased and N remobilization fluxes were enhanced from source organs (leaves and stem) toward sink organs (flowers). Profuse branching and late senescing varieties would be of interest for further selection programs under high sowing densities.

Background In cereal crops such as wheat, an optimal timing of developmental transitions is required to maximize grain yield. Many of these developmental changes are precisely regulated by changes in the duration, intensity or quality of light. Phytochromes are dimeric photoreceptors that absorb light maximally in the red and far-red wavelengths and induce large-scale transcriptional changes in response to variation in light quality. In wheat, PHYC is required for early flowering under long days. However, it is currently unknown whether this function requires the presence of PHYB. In this study, we characterized the role of PHYB in wheat development and used RNA-seq to analyze and compare the transcriptomes of phyB-null and phyC-null TILLING mutants. Results Under long-day photoperiods, phyB-null plants exhibit a severe delay in flowering comparable to the delay observed in phyC-null plants. These results demonstrate that both genes are required for the induction of wheat flowering under long days. Using replicated RNA-seq studies we identified 82 genes that are significantly up or down regulated in both the phyB-null and phyC-null mutant relative to their respective wild-type controls. Among these genes are several well-characterized positive regulators of flowering, including PPD1, FT1 and VRN1. Eight-fold more genes were differentially regulated only in the phyB-null mutant (2202) than only in the phyC-null mutant (261). The PHYB-regulated genes were enriched in components of the auxin, gibberellin and brassinosteroid biosynthesis and signaling pathways, and in transcription factors with putative roles in regulating vegetative development and shade-avoidance responses. Several genes involved in abiotic stress tolerance pathways were also found to be regulated by PHYB. Conclusions PHYB and PHYC are both required for the photoperiodic induction of wheat flowering, whereas PHYB alone regulates a large number of genes involved in hormone biosynthesis and signaling, shade-avoidance response, and abiotic stress tolerance. Our analysis provides a comprehensive overview of the PHYB- and PHYC-mediated transcriptional changes during light signaling, and an initial step towards the dissection of this regulatory gene network in wheat. This further dissection will be required to explore the individual phytochrome-mediated developmental responses and to evaluate their potential to improve wheat adaptation to changing environments. Electronic supplementary material The online version of this article (doi:10.1186/s12870-016-0831-3) contains supplementary material, which is available to authorized users.

Plant sterols and steroid hormones, the brassinosteroids (BRs), are compounds that exert a wide range of biological activities. They are essential for plant growth, reproduction, and responses to various abiotic and biotic stresses. Given the importance of sterols and BRs in these processes, engineering their biosynthetic and signaling pathways offers exciting potentials for enhancing crop yield. In this review, we focus on how alterations in components of sterol and BR metabolism and signaling or application of exogenous steroids and steroid inhibitors affect traits of agronomic importance. We also discuss areas for future research and identify the fine-tuning modulation of endogenous BR content as a promising strategy for crop improvement.

Different genotypes often exhibit opposite plastic responses in the timing of the onset of flowering with increasing plant density. In experimental studies, selection for accelerated flowering is generally found. By contrast, game theoretical studies predict that there should be selection for delayed flowering when competition increases. Combining different optimality criteria, the conditions under which accelerated or delayed flowering in response to density would be selected for are analysed with a logistic growth simulation model. To maximize seed production at the whole-stand level (simple optimization), selection should lead to accelerated flowering at high plant density, unless very short growing seasons select for similar onset of flowering at all densities. By contrast, selection of relative individual fitness will lead to delayed flowering when season length is long and/or growth rates are high. These different results give a potential explanation for the observed differences in direction of the plastic responses within and between species, including homeostasis, as a result of the effect of the variation in season length on the benefits of delayed flowering. This suggests that limited plasticity can evolve without the costs and limits that are currently thought to constrain the evolution of plasticity.

Although plants are sessile organisms, they can modulate their phenotype so as to cope with environmental stresses such as herbivore attack and competition with neighbouring plants. Plant-produced volatile compounds mediate various aspects of plant defence. The emission of volatiles has costs and benefits. Research on the role of plant volatiles in defence has focused primarily on the responses of individual plants. However, in nature, plants rarely occur as isolated individuals but are members of plant communities where they compete for resources and exchange information with other plants. In this review, we address the effects of neighbouring plants on plant volatile-mediated defences. We will outline the various roles of volatile compounds in the interactions between plants and other organisms, address the mechanisms of plant neighbour perception in plant communities, and discuss how neighbour detection and volatile signalling are interconnected. Finally, we will outline the most urgent questions to be addressed in the future. Fil: Pierik, Ronald. Utrecht University. Institute of Environmental Biology. Plant Ecophysiology; Países Bajos Fil: Ballare, Carlos Luis. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico La Plata. Instituto de Investigaciones Biotecnológicas - Instituto Tecnológico Chascomús. Instituto de Investigaciones Biotecnológicas (sede Chascomús); Argentina Fil: Dicke, Marcel. Wageningen University. Laboratory of Entomology; Países Bajos

J1: Front Plant Sci M1 - Journal Article Control of flowering in the perennial model, the woodland strawberry (Fragaria vesca L.), involves distinct molecular mechanisms that result in contrasting photoperiodic flowering responses and growth cycles in different accessions. The F. vesca homolog of TERMINAL FLOWER1 (FvTFL1) functions as a key floral repressor that causes short-day (SD) requirement of flowering and seasonal flowering habit in the SD strawberry. In contrast, perpetual flowering F. vesca accessions lacking functional FvTFL1 show FLOWERING LOCUS T (FvFT1)-dependent early flowering specifically under long-days (LD). We show here that the end-of-day far-red (FR) and blue (B) light activate the expression of FvFT1 and the F. vesca homolog of SUPPRESSOR OF THE OVEREXPRESSION OF CONSTANS (FvSOC1) in both SD and LD strawberries, whereas low expression levels are detected in red (R) and SD treatments. By using transgenic lines, we demonstrate that FvFT1 advances flowering under FR and B treatments compared to R and SD treatments in the LD strawberry, and that FvSOC1 is specifically needed for the B light response. In the SD strawberry, flowering responses to these light quality treatments are reversed due to up-regulation of the floral repressor FvTFL1 in parallel with FvFT1 and FvSOC1. Our data highlights the central role of FvFT1 in the light quality dependent flower induction in the LD strawberry and demonstrates that FvTFL1 reverses not only photoperiodic requirements but also light quality effects on flower induction in the SD strawberry.

Competition and parasitism are two important selective forces that shape life-histories, migration rates and population dynamics. Recently, it has been shown in various pathosystems that parasites can modify intraspecific competition, thus generating an indirect cost of parasitism. Here, we investigated if this phenomenon was present in a plant-potyvirus system using two viruses of different virulence (Tobacco etch virus and Turnip mosaic virus). Moreover, we asked if parasitism interacted with the shade avoidance syndrome, the plant-specific phenotypic plasticity in response to intraspecific competition. Our results indicate that the modification of intraspecific competition by parasitism is not present in the Nicotiana benthamiana – potyvirus system and suggests that this phenomenon is not universal but depends on the peculiarities of each pathosystem. However, whereas the healthy N. benthamiana presented a clear shade avoidance syndrome, this phenotypic plasticity totally disappeared when the plants were infected with TEV and TuMV, very likely resulting in a fitness loss and being another form of indirect cost of parasitism. This result suggests that the suppression or the alteration of adaptive phenotypic plasticity might be a component of virulence that is often overlooked., This work has been supported by grants BFU2009-06993 (Ministerio de Ciencia e Innovacio´ n, Spain) and PROMETEO2010-019 (Generalitat Valenciana) (S.F.E.) and a CSIC JAE-Doc contract (S.B.). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Plants can respond quickly and profoundly to detrimental changes in their environment. For example, Arabidopsis thaliana can induce an upward leaf movement response through differential petiole growth (hyponastic growth) to outgrow complete submergence. This response is induced by accumulation of the phytohormone ethylene in the plant. Currently, only limited information is available on how this response is molecularly controlled. In this study, we utilized quantitative trait loci (QTL) analysis of natural genetic variation among Arabidopsis accessions to isolate novel factors controlling constitutive petiole angles and ethylene-induced hyponastic growth. Analysis of mutants in various backgrounds and complementation analysis of naturally occurring mutant accessions provided evidence that the leucin-rich repeat receptor-like Ser/Thr kinase gene, ERECTA, controls ethylene-induced hyponastic growth. Moreover, ERECTA controls leaf positioning in the absence of ethylene treatment. Our data demonstrate that this is not due to altered ethylene production or sensitivity.

The current knowledge of light quality effects on plant morphogenesis and development represents a new era of understanding on how plant communities perceive and adjust to available resources. The most important consequences of light quality cues, often mediated by decreasing in red far-red ratios with respect to the spectral composition of incident sunlight radiation, affecting weed-crop interaction are the increased plant height and shoot to root ratio in anticipation of competition by light quantity, water or nutrients. Although the concepts related to light quality have been extensively studied and several basic process of this phenomenon are well known, little applications of photomorphogenic signaling currently are related to agricultural problems or weed management. The objectives of this review are to describe how light quality change can be a triggering factor of interspecific interference responses, to analyze how this phenomenon can be used to predict weed interference, to reevaluate the critical periods of interference concept, and to discuss its potential contribution towards developing more weed competitive crop varieties. Knowledge on light quality responses involved in plant sensing of interspecific competition could be used to identify red/far-red threshold values, indicating when weed control should be started. Light quality alterations by weeds can affect grain crop development mainly in high yielding fields. Unlike the traditional concept or the critical period of competition, light quality mediated interference implies that the critical period for weed control could start before the effects of direct resource (water, nutrients and available light) limitation actually occur. The variability in light quality responses among crop genotypes and the identification of mutants insensitive to light quality effects indicate that this characteristic can be selected or modified to develop cultivars with enhanced interspecific interference ability. Knowledge on light quality-elicited responses represents a new possibility to understand the underlying biology of interspecific interference, and could be used in the development of new weed management technologies. O atual conhecimento sobre os efeitos da qualidade da luz representa uma nova era da compreensão dos processos envolvidos na percepção e ajuste das plantas em função dos recursos disponíveis no ambiente. As consequências mais importantes da diminuição da qualidade de luz, que se relaciona principalmente com a diminuição da relação vermelho/vermelho-extremo, que ocorrem nas interações entre culturas e plantas daninhas são o aumento da estatura e da relação parte aérea/raiz. Essas alterações são tentativas de antecipar os efeitos da concorrência futura por quantidade de luz, água ou nutrientes. Embora vários conhecimentos básicos relacionados com a qualidade da luz tenham sido desenvolvidos há muito tempo, poucas aplicações da sinalização fotomorfogênica têm sido implementadas em processos relacionados com a agricultura e, principalmente, com o manejo de plantas daninhas. Os objetivos deste trabalho foram: descrever a diminuição da qualidade da luz como um fator desencadeante da competição interespecífica; analisar como a qualidade da luz pode ser usada para predizer o início da interferência das plantas daninhas; reavaliar os períodos críticos de interferência; e discutir o potencial de desenvolvimento de cultivares mais competitivas com as plantas daninhas. O conhecimento dos fatores envolvidos na qualidade da luz pode ser utilizado para identificação de uma relação da radiação vermelho/vermelho-distante que indique o início da competição interespecífica e possa ser usada com um nível de dano referencial, indicando quando o controle das plantas daninhas deve ser iniciado. Os efeitos da baixa qualidade de luz podem limitar o desenvolvimento das culturas, sobretudo em situações de lavouras com elevados níveis potenciais de rendimento de grãos. Nessas situações, o período crítico para controle de plantas daninhas deve ser considerado antes de se iniciarem os efeitos da limitação dos recursos de água, nutrientes e quantidade de luz, que representa o tradicional período crítico de competição. A existência de variabilidade de genótipos das culturas à qualidade da luz e a identificação de mutantes insensíveis aos efeitos da baixa qualidade da luz indicam que essa característica pode ser selecionada ou transformada para o desenvolvimento de cultivares com maior capacidade de competição com as plantas daninhas. Os conhecimentos relacionados com a qualidade da luz representam uma nova possibilidade de compreender a biologia da competição interespecífica e poderão ser utilizados no desenvolvimento de novas tecnologias para o manejo de plantas daninhas.

O atual conhecimento sobre os efeitos da qualidade da luz representa uma nova era da compreensão dos processos envolvidos na percepção e ajuste das plantas em função dos recursos disponíveis no ambiente. As consequências mais importantes da diminuição da qualidade de luz, que se relaciona principalmente com a diminuição da relação vermelho/ vermelho-extremo, que ocorrem nas interações entre culturas e plantas daninhas são o aumento da estatura e da relação parte aérea/raiz. Essas alterações são tentativas de antecipar os efeitos da concorrência futura por quantidade de luz, água ou nutrientes. Embora vários conhecimentos básicos relacionados com a qualidade da luz tenham sido desenvolvidos há muito tempo, poucas aplicações da sinalização fotomorfogênica têm sido implementadas em processos relacionados com a agricultura e, principalmente, com o manejo de plantas daninhas. Os objetivos deste trabalho foram: descrever a diminuição da qualidade da luz como um fator desencadeante da competição interespecífica; analisar como a qualidade da luz pode ser usada para predizer o início da interferência das plantas daninhas; reavaliar os períodos críticos de interferência; e discutir o potencial de desenvolvimento de cultivares mais competitivas com as plantas daninhas. O conhecimento dos fatores envolvidos na qualidade da luz pode ser utilizado para identificação de uma relação da radiação vermelho/ vermelho-distante que indique o início da competição interespecífica e possa ser usada com um nível de dano referencial, indicando quando o controle das plantas daninhas deve ser iniciado. Os efeitos da baixa qualidade de luz podem limitar o desenvolvimento das culturas, sobretudo em situações de lavouras com elevados níveis potenciais de rendimento de grãos. Nessas situações, o período crítico para controle de plantas daninhas deve ser considerado antes de se iniciarem os efeitos da limitação dos recursos de água, nutrientes e quantidade de luz, que representa o tradicional período crítico de competição. A existência de variabilidade de genótipos das culturas à qualidade da luz e a identificação de mutantes insensíveis aos efeitos da baixa qualidade da luz indicam que essa característica pode ser selecionada ou transformada para o desenvolvimento de cultivares com maior capacidade de competição com as plantas daninhas. Os conhecimentos relacionados com a qualidade da luz representam uma nova possibilidade de compreender a biologia da competição interespecífica e poderão ser utilizados no desenvolvimento de novas tecnologias para o manejo de plantas daninhas. The current knowledge of light quality effects on plant morphogenesis and development represents a new era of understanding on how plant communities perceive and adjust to available resources. The most important consequences of light quality cues, often mediated by decreasing in red far-red ratios with respect to the spectral composition of incident sunlight radiation, affecting weed-crop interaction are the increased plant height and shoot to root ratio in anticipation of competition by light quantity, water or nutrients. Although the concepts related to light quality have been extensively studied and several basic process of this phenomenon are well known, little applications of photomorphogenic signaling currently are related to agricultural problems or weed management. The objectives of this review are to describe how light quality change can be a triggering factor of interspecific interference responses, to analyze how this phenomenon can be used to predict weed interference, to reevaluate the critical periods of interference concept, and to discuss its potential contribution towards developing more weed competitive crop varieties. Knowledge on light quality responses involved in plant sensing of interspecific competition could be used to identify red/far-red threshold values, indicating when weed control should be started. Light quality alterations by weeds can affect grain crop development mainly in high yielding fields. Unlike the traditional concept or the critical period of competition, light quality mediated interference implies that the critical period for weed control could start before the effects of direct resource (water, nutrients and available light) limitation actually occur. The variability in light quality responses among crop genotypes and the identification of mutants insensitive to light quality effects indicate that this characteristic can be selected or modified to develop cultivars with enhanced interspecific interference ability. Knowledge on light quality-elicited responses represents a new possibility to understand the underlying biology of interspecific interference, and could be used in the development of new weed management technologies.

Upward leaf movement (hyponastic growth) is frequently observed in response to changing environmental conditions and can be induced by the phytohormone ethylene. Hyponasty results from differential growth (i.e. enhanced cell elongation at the proximal abaxial side of the petiole relative to the adaxial side). Here, we characterize Enhanced Hyponasty-d, an activation-tagged Arabidopsis (Arabidopsis thaliana) line with exaggerated hyponasty. This phenotype is associated with overexpression of the mitotic cyclin CYCLINA2;1 (CYCA2;1), which hints at a role for cell divisions in regulating hyponasty. Indeed, mathematical analysis suggested that the observed changes in abaxial cell elongation rates during ethylene treatment should result in a larger hyponastic amplitude than observed, unless a decrease in cell proliferation rate at the proximal abaxial side of the petiole relative to the adaxial side was implemented. Our model predicts that when this differential proliferation mechanism is disrupted by either ectopic overexpression or mutation of CYCA2;1, the hyponastic growth response becomes exaggerated. This is in accordance with experimental observations on CYCA2;1 overexpression lines and cyca2;1 knockouts. We therefore propose a bipartite mechanism controlling leaf movement: ethylene induces longitudinal cell expansion in the abaxial petiole epidermis to induce hyponasty and simultaneously affects its amplitude by controlling cell proliferation through CYCA2;1. Further corroborating the model, we found that ethylene treatment results in transcriptional down-regulation of A2-type CYCLINs and propose that this, and possibly other regulatory mechanisms affecting CYCA2;1, may contribute to this attenuation of hyponastic growth.

Shade-avoidance is a major adaptive response of plants, and is usually considered to be controlled by phytochromes through the perception of changes in the red:far red light ratio. However, few studies on the effects of blue light (BL) and of light intensity [photosynthetically active radiation (PAR)] on light-grown plants have been conducted, especially concerning changes in PAR at constant BL. The objective here was to quantify the photocontrol of aerial morphogenesis by BL and PAR. Experiments were conducted varying BL and PAR independently, with three BL levels (4, 38, and 83 micromol m(-2) s(-1)) at constant PAR (300 micromol m(-2) s(-1)) and three PAR levels (338, 705, and 163 micromol m(-2) s(-1)) at constant BL (36 micromol m(-2) s(-1)). Effects on morphogenetic processes were analysed as quantitative modulations of ontogenic trends and response curves were produced. White clover (Trifolium repens L.) was used, as it is a typical shade-avoider displaying the whole syndrome of shade-avoidance in a purely vegetative stage. Morphological responses were strongly controlled by both BL and PAR changes, through antagonist effects on leaf appearance rate and additive effects on petiole elongation. All the other responses appeared to be the indirect consequences of changes in the leaf appearance rates. BL acted as a light signal for plant morphogenesis. However, the PAR control probably implicates two distinct mechanisms, such as a trophic effect and a signal. Both PAR and BL actions involved organ-specific differences, which are central in the control of the shade-avoidance responses.

Radiation substantially affects leaf initiation rate (LIR), a key variable for plant growth, by influencing the heat budget and therefore the temperature of the shoot apical meristem. The photosynthetically active component of solar radiation (photosynthetic photon flux density; PPFD) is critical for plant growth and when at shade to moderate levels may also influence LIR via limited photosynthate availability. Cucumber and tomato plants were subjected to different PPFDs (2.5–13.2 mol m–2 day–1) and then LIR, carbohydrate content and diel net CO2 uptake of the apical bud were quantified. LIR showed saturating response to increasing PPFD in both species. In this PPFD range, LIR was reduced by 20% in cucumber and by 40% in tomato plants. Carbohydrate content and dark respiration were substantially reduced at low PPFD. LIR may be considered as an adaptive trait of plants to low light levels, which is likely to be determined by the local photosynthate availability. In tomato and cucumber plants, LIR can be markedly reduced at low PPFD in plant production systems at high latitudes, suggesting that models solely based on thermal time may not precisely predict LIR at low PPFD.