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Background and Aims Leaf elongation is vital for productivity of Poaceae species, influenced by atmospheric CO2 concentration ([CO2]) and climate-induced water availability changes. Although [CO2] mitigates the effects of drought on reducing transpiration per unit leaf area, it also increases total leaf area and water use. These complex interactions associated with leaf growth pose challenges in anticipating climate change effects. This study aims to assess [CO2] effects on leaf growth response to drought in perennial ryegrass (Lolium perenne), tall fescue (Festuca arundinacea) and wheat (Triticum aestivum). Methods Plants were cultivated in growth chambers with [CO2] at 200 or 800 ppm. At leaf six to seven unfolding, half of the plants were subjected to severe drought treatment. Leaf elongation rate (LER) was measured daily, whereas plant transpiration was continuously recorded gravimetrically. Additionally, water-soluble carbohydrate (WSC) content along with water and osmotic potentials in the leaf growing zone were measured at drought onset, mid-drought and leaf growth cessation. Key Results Elevated [CO2] mitigated drought impacts on LER and delayed growth cessation across species. A positive correlation between LER and soil relative water content (SRWC) was observed. At the same SRWC, perennial grasses exhibited a higher LER with elevated [CO2], probably due to enhanced stomatal regulation. Despite stomatal closure and WSC accumulation, CO2 did not influence nighttime water potential or osmotic potential. The marked increase in leaf area across species resulted in similar (wheat and tall fescue) or higher (ryegrass) total water use by the end of the experiment, under both watered and unwatered conditions. Conclusions Elevated [CO2] mitigates the adverse effects of drought on leaf elongation in three Poaceae species, due to its impact on plant transpiration. Overall, these findings provide valuable insights into CO2 and drought interactions that may help anticipate plant responses to climate change. [ABSTRACT FROM AUTHOR]

Foliar development involves successive phases of cell proliferation and expansion that determine the final leaf size, and is characterized by an early burst of reactive oxygen species generated in the photosynthetic electron transport chain (PETC). Introduction of the alternative PETC acceptor flavodoxin in tobacco chloroplasts led to a reduction in leaf size associated to lower cell expansion, without affecting cell number per leaf. Proteomic analysis showed that the biogenesis of the PETC proceeded stepwise in wild-type leaves, with accumulation of light-harvesting proteins preceding that of electron transport components, which might explain the increased energy and electron transfer to oxygen and reactive oxygen species build-up at this stage. Flavodoxin expression did not affect biogenesis of the PETC but prevented hydroperoxide formation through its function as electron sink. Mature leaves from flavodoxin-expressing plants were shown to contain higher levels of transcripts encoding components of the proteasome, a key negative modulator of organ size. Proteome profiling revealed that this differential accumulation was initiated during expansion and led to increased proteasomal activity, whereas a proteasome inhibitor reverted the flavodoxin-dependent size phenotype. Cells expressing plastid-targeted flavodoxin displayed lower endoreduplication, also associated to decreased organ size. These results provide novel insights into the regulation of leaf growth by chloroplast-generated redox signals, and highlight the potential of alternative electron shuttles to investigate the link(s) between photosynthesis and plant development. [ABSTRACT FROM AUTHOR]

The leaf is a determinate organ with a final size under genetic control. Numerous factors that regulate the final leaf size have been identified in Arabidopsis thaliana ; although most of these factors play their roles during the growth of leaf primordia, much less is known about leaf initiation and its effects on the final leaf size. In this study, we characterized oligocellula6-D (oli6-D), a semidominant mutant of A. thaliana with smaller leaves than the wild type (WT) due to its reduced leaf cell numbers. A time-course analysis showed that oli6-D had approximately 50% fewer leaf cells even immediately after leaf initiation; this difference was maintained throughout leaf development. Next-generation sequencing showed that oli6-D had chromosomal duplications involving 2-kb and 3-Mb regions of chromosomes 2 and 4, respectively. Several duplicated genes examined had approximately 2-fold higher expression levels, and at least one gene acquired a new intron/exon structure due to a chromosome fusion event. oli6-D showed reduced auxin responses in leaf primordia, primary roots and embryos, as well as reduced apical dominance and partial auxin-resistant root growth. CRISPR-associated protein-9-mediated genome editing enabled the removal of a 3-Mb duplicated segment, the largest targeted deletion in plants thus far. As a result, oli6-D restored the WT leaf phenotypes, demonstrating that oli6-D is a gain-of-function mutant. Our results suggest a new regulatory point of leaf size determination that functions at a very early stage of leaf development and is negatively regulated by one or more genes located in the duplicated chromosomal segments. [ABSTRACT FROM AUTHOR]

Background and Aims Intra- and transgenerational plasticity may provide substantial phenotypic variation to cope with environmental change. Since assessing the unique contribution of the maternal environment to the offspring phenotype is challenging in perennial, outcrossing plants, little is known about the evolutionary and ecological implications of transgenerational plasticity and its persistence over the life cycle in these species. We evaluated how intra- and transgenerational plasticity interplay to shape the adaptive responses to drought in two perennial Mediterranean shrubs. Methods We used a novel common garden approach that reduced within-family genetic variation in both the maternal and offspring generations by growing the same maternal individual in two contrasting watering environments, well-watered and drought, in consecutive years. We then assessed phenotypic differences at the reproductive stage between offspring reciprocally grown in the same environments. Key Results Maternal drought had an effect on offspring performance only in Helianthemum squamatum. Offspring of drought-stressed plants showed more inflorescences, less sclerophyllous leaves and higher growth rates in both watering conditions, and heavier seeds under drought, than offspring of well-watered maternal plants. Maternal drought also induced similar plasticity patterns across maternal families, showing a general increase in seed mass in response to offspring drought, a pattern not observed in the offspring of well-watered plants. In contrast, both species expressed immediate adaptive plasticity, and the magnitude of intragenerational plasticity was larger than the transgenerational plastic responses. Conclusions Our results highlight that adaptive effects associated with maternal drought can persist beyond the seedling stage and provide evidence of species-level variation in the expression of transgenerational plasticity. Such differences between co-occurring Mediterranean species in the prevalence of this form of non-genetic inheritance may result in differential vulnerability to climate change. [ABSTRACT FROM AUTHOR]

Forest trees adopt effective strategies to optimize nitrogen (N) use through internal N recycling. In the context of more recurrent environmental stresses due to climate change, the question remains of whether increased frequency of drought or defoliation threatens this internal N recycling strategy. We submitted 8-year-old beech trees to 2 years of either severe drought (Dro) or manual defoliation (Def) to create a state of N starvation. At the end of the second year before leaf senescence, we labeled the foliage of the Dro and Def trees, as well as that of control (Co) trees, with 15N-urea. Leaf N resorption, winter tree N storage (total N, 15N, amino acids, soluble proteins) and N remobilization in spring were evaluated for the three treatments. Defoliation and drought did not significantly impact foliar N resorption or N concentrations in organs in winter. Total N amounts in Def tree remained close to those in Co tree, but winter N was stored more in the branches than in the trunk and roots. Total N amount in Dro trees was drastically reduced (−55%), especially at the trunk level, but soluble protein concentrations increased in the trunk and fine roots compared with Co trees. During spring, 15N was mobilized from the trunk, branches and twigs of both Co and Def trees to support leaf growth. It was only provided through twig 15N remobilization in the Dro trees, thus resulting in extremely reduced Dro leaf N amounts. Our results suggest that stress-induced changes occur in N metabolism but with varying severity depending on the constraints: within-tree 15N transport and storage strategy changed in response to defoliation, whereas a soil water deficit induced a drastic reduction of the N amounts in all the tree organs. Consequently, N dysfunction could be involved in drought-induced beech tree mortality under the future climate. [ABSTRACT FROM AUTHOR]

C4 photosynthesis outperforms C3 photosynthesis in natural ecosystems by maintaining a high photosynthetic rate and affording higher water-use and nitrogen-use efficiencies. C4 plants can survive in environments with poor living conditions, such as high temperatures and arid regions, and will be crucial to ecological and agricultural security in the face of global climate change in the future. However, the genetic architecture of C4 photosynthesis remains largely unclear, especially the genetic regulation of C4 Kranz anatomy. Haloxylon ammodendron is an important afforestation tree species and a valuable C4 wood plant in the desert region. The unique characteristic of H. ammodendron is that, during the seedling stage, it utilizes C3 photosynthesis, while in mature assimilating shoots (maAS), it switches to the C4 pathway. This makes an exceptional opportunity for studying the development of the C4 Kranz anatomy and metabolic pathways within individual plants (identical genome). To provide broader insight into the regulation of Kranz anatomy and non-Kranz leaves of the C4 plant H. ammodendron , carbon isotope values, anatomical sections and transcriptome analyses were used to better understand the molecular and cellular processes related to the development of C4 Kranz anatomy. This study revealed that H. ammodendron conducts C3 in the cotyledon before it switches to C4 in AS. However, the switching requires a developmental process. Stable carbon isotope discrimination measurements on three different developmental stages showed that young AS have a C3-like δ13C even though C4 Kranz anatomy is found, which is inconsistent with the anatomical findings. A C4-like δ13C can be measured in AS until they are mature. The expression analysis of C4 key genes also showed that the maAS exhibited higher expression than the young AS. In addition, many genes that may be related to the development of Kranz anatomy were screened. Comparison of gene expression patterns with respect to anatomy during leaf ontogeny provided insight into the genetic features of Kranz anatomy. This study helps with our understanding of the development of Kranz anatomy and provides future directions for studies on key C4 regulatory genes. [ABSTRACT FROM AUTHOR]

The growth of fruit trees depends on the nitrogen (N) remobilization in mature tissues and N acquisition from the soil. However, in evergreen mature citrus (Citrus reticulata Blanco) leaves, proteins with N storage functions and hub molecules involved in driving N remobilization remain largely unknown. Here, we combined proteome and physiological analyses to characterize the spatiotemporal mechanisms of growth of new leaves and storage protein degradation in mature leaves of citrus trees exposed to low-N and high-N fertilization in the field. Results show that the growth of new leaves is driven by remobilization of stored reserves, rather than N uptake by the roots. In this context, proline and arginine in mature leaves acted as N sources supporting the growth of new leaves in spring. Time-series analyses with gel electrophoresis and proteome analysis indicated that the mature autumn shoot leaves are probably the sites of storage protein synthesis, while the aspartic endopeptidase protein is related to the degradation of storage proteins in mature citrus leaves. Furthermore, bioinformatic analysis based on protein–protein interactions indicated that glutamate synthetase and ATP-citrate synthetase are hub proteins in N remobilization from mature citrus leaves. These results provide strong physiological data for seasonal optimization of N fertilizer application in citrus orchards. [ABSTRACT FROM AUTHOR]

Twenty-five years ago, a seminal paper demonstrated that warm temperatures increase auxin levels to promote hypocotyl growth in Arabidopsis thaliana. Here we highlight recent advances in auxin-mediated thermomorphogenesis and identify unanswered questions. In the warmth, PHYTOCHROME INTERACTING FACTOR 4 (PIF4) and PIF7 bind the YUCCA8 gene promoter and, in concert with histone modifications, enhance its expression to increase auxin synthesis in the cotyledons. Once transported to the hypocotyl, auxin promotes cell elongation. The meta-analysis of expression of auxin-related genes in seedlings exposed to temperatures ranging from cold to hot shows complex patterns of response. Changes in auxin only partially account for these responses. The expression of many SMALL AUXIN UP RNA (SAUR) genes reaches a maximum in the warmth, decreasing towards both temperature extremes in correlation with the rate of hypocotyl growth. Warm temperatures enhance primary root growth, the response requires auxin, and the hormone levels increase in the root tip but the impacts on cell division and cell expansion are not clear. A deeper understanding of auxin-mediated temperature control of plant architecture is necessary to face the challenge of global warming. [ABSTRACT FROM AUTHOR]

Triose phosphate utilization (TPU) is a biochemical process indicating carbon sink–source (im)balance within leaves. When TPU limits leaf photosynthesis, photorespiration-associated amino acid exports probably provide an additional carbon outlet and increase leaf CO2 uptake. However, whether TPU is modulated by whole-plant sink–source relations and nitrogen (N) budgets remains unclear. We address this question by model analyses of gas-exchange data measured on leaves at three growth stages of rice plants grown at two N levels. Sink–source ratio was manipulated by panicle pruning, by using yellower-leaf variant genotypes, and by measuring photosynthesis on adaxial and abaxial leaf sides. Across all these treatments, higher leaf N content resulted in the occurrence of TPU limitation at lower intercellular CO2 concentrations. Photorespiration-associated amino acid export was greater in high-N leaves, but was smaller in yellower-leaf genotypes, panicle-pruned plants, and for abaxial measurement. The feedback inhibition of panicle pruning on rates of TPU was not always observed, presumably because panicle pruning blocked N remobilization from leaves to grains and the increased leaf N content masked feedback inhibition. The leaf-level TPU limitation was thus modulated by whole-plant sink–source relations and N budgets during rice grain filling, suggesting a close link between within-leaf and whole-plant sink limitations. [ABSTRACT FROM AUTHOR]

Background and Aims Plasticity of leaf growth and photosynthesis is an important strategy of plants to adapt to shading stress; however, their strategy of leaf development to achieve a simultaneous increase in leaf area and photosynthesis under shading remains unknown. Methods In the present study, a pot experiment was conducted using three rapeseed genotypes of Huayouza 50 (HYZ50), Zhongshuang 11 (ZS11) and Huayouza 62 (HYZ62), and the responses of plant growth, leaf morphoanatomical traits, cell wall composition and photosynthesis to shading were investigated. Key Results Shading significantly increased leaf area per plant (LA plant) in all genotypes, but the increase in HYZ62 was greater than that in HYZ50 and ZS11. The greater increment of LA plant in HYZ62 was related to the larger decrease in leaf mass per area (LMA) and leaf density (LD), which were in turn related to less densely packed mesophyll cells and thinner cell walls (T cw). Moreover, shading significantly increased photosynthesis in HYZ62 but significantly decreased it in HYZ50. The enhanced photosynthesis in HYZ62 was related to increased mesophyll conductance (g m) due primarily to thinner cell walls. Conclusions The data presented indicate that the different plasticity of mesophyll cell density, cell wall thickness and cell wall composition in response to shading can dramatically affect leaf growth and photosynthesis. [ABSTRACT FROM AUTHOR]

Background and Aims Drought events have devasting impacts on grape berry production. The aim of this study was to investigate berry growth in the context of leaf stomatal closure under progressive drought stress. Methods Potted grapevine plants (varieties 'Syrah' and 'Cabernet Sauvignon') were evaluated at pre-verasion (30–45 d after anthesis, DAA) and post-veraison (90–107 DAA). Berry diameter, berry absolute growth rate (AGR), leaf stomatal conductance (G s) at midday, plant water potential at predawn and midday (ΨPD and ΨMD, respectively), and soil relative water content were measured repeatedly. The ΨPD-threshold of 90 % loss in stomatal conductance (G s10, i.e. complete stomatal closure) was determined. Data were related to plant dehydration phases I, II and III with corresponding boundaries Θ1 and Θ2, using the water potential curve method. Key Results At pre-veraison, berry AGR declined together with leaf G s in response to soil drying in both varieties. Berry AGR transitioned from positive to negative (shrinkage) values when leaf G s approached zero. The G s10-threshold was −0.81 MPa in 'Syrah' and −0.74 MPa in 'Cabernet Sauvignon' and was linked to boundary Θ1. At post-veraison, berry AGR was negligible and negative AGR values were not intensified by increasing drought stress in either variety. Conclusion Leaf complete stomatal closure under progressive drought stress coincides with cessation of berry growth followed by shrinkage at pre-veraison (growth stage 1). [ABSTRACT FROM AUTHOR]

Nitrogen (N) is an essential macronutrient for plants, and its remobilization is key for adaptation to deficiency stress. However, there is limited understanding of the regulatory mechanisms of N remobilization in the important crop species Brassica napus (oilseed rape). Here, we report the identification of a transcription factor, BnaA9.WRKY47 , that is induced by N starvation in a canola variety. At the seedling stage, BnaA9.WRKY47 -overexpressing (OE) lines displayed earlier senescence of older leaves and preferential growth of juvenile leaves compared to the wild type under N starvation. At the field scale, the seed yield was significantly increased in the BnaA9.WRKY47 -OE lines compared with the wild type when grown under N deficiency conditions and, conversely, it was reduced in BnaA9.WRKY47 -knockout mutants. Biochemical analyses demonstrated that BnaA9.WRKY47 directly activates BnaC7.SGR1 to accelerate senescence of older leaves. In line with leaf senescence, the concentration of amino acids in the older leaves of the OE lines was elevated, and the proportion of plant N that they contained was reduced. This was associated with BnaA9.WRKY47 activating the amino acid permease BnaA9.AAP1 and the nitrate transporter BnaA2.NRT1.7. Thus, the expression of BnaA9.WRKY47 efficiently facilitated N remobilization from older to younger leaves or to seeds. Taken together, our results demonstrate that BnaA9.WRKY47 up-regulates the expression of BnaC7.SGR1 , BnaA2.NRT1.7 , and BnaA9AAP1 , thus promoting the remobilization of N in B. napus under starvation conditions. [ABSTRACT FROM AUTHOR]

Plant growth is a complex process affected by a multitude of genetic and environmental factors and their interactions. To identify genetic factors influencing plant performance under different environmental conditions, vegetative growth was assessed in Arabidopsis thaliana cultivated under constant or fluctuating light intensities, using high-throughput phenotyping and genome-wide association studies. Daily automated non-invasive phenotyping of a collection of 382 Arabidopsis accessions provided growth data during developmental progression under different light regimes at high temporal resolution. Quantitative trait loci (QTL) for projected leaf area, relative growth rate, and PSII operating efficiency detected under the two light regimes were predominantly condition-specific and displayed distinct temporal activity patterns, with active phases ranging from 2 d to 9 d. Eighteen protein-coding genes and one miRNA gene were identified as potential candidate genes at 10 QTL regions consistently found under both light regimes. Expression patterns of three candidate genes affecting projected leaf area were analysed in time-series experiments in accessions with contrasting vegetative leaf growth. These observations highlight the importance of considering both environmental and temporal patterns of QTL/allele actions and emphasize the need for detailed time-resolved analyses under diverse well-defined environmental conditions to effectively unravel the complex and stage-specific contributions of genes affecting plant growth processes. [ABSTRACT FROM AUTHOR]

Studying intraspecific variation in multistress responses is central for predicting and managing the population dynamics of wild plant species under rapid global change. Yet, it remains a challenging goal in this field to integrate knowledge on the complex biochemical underpinnings for the targeted 'non-model' species. Here, we studied divergence in combined drought and heat responses among Northern and Southern European populations of the dune plant Cakile maritima , by combining comprehensive plant phenotyping with metabolic profiling via FT-ICR-MS and UPLC-TQ-MS/MS. We observed pronounced constitutive divergence in growth phenology, leaf functional traits, and defence chemistry (glucosinolates and alkaloids) among population origins. Most importantly, the magnitude of growth reduction under drought was partly weaker in southern plants and associated with divergence in plastic growth responses (leaf abscission) and the modulation of primary and specialized metabolites with known central functions not only in plant abiotic but also in biotic stress responses. Our study indicates that divergent selection has shaped the constitutive and drought-/heat-induced expression of numerous morphological and biochemical functional traits to mediate higher abiotic stress resistance in southern Cakile populations, and highlights that metabolomics can be a powerful tool to explore the underlying mechanisms of local adaptation in 'non-model' species. [ABSTRACT FROM AUTHOR]

C3 and C4 grasses directly and indirectly provide the vast majority of calories to the human diet, yet our understanding of the molecular mechanisms driving photosynthetic productivity in grasses is largely unexplored. Ground meristem cells divide to form mesophyll or vascular initial cells early in leaf development in C3 and C4 grasses. Here we define a genetic circuit composed of SHORT ROOT (SHR), INDETERMINATE DOMAIN (IDD), and PIN-FORMED (PIN) family members that specifies vascular identify and ground cell proliferation in leaves of both C3 and C4 grasses. Ectopic expression and loss-of-function mutant studies of SHR paralogs in the C3 plant Oryza sativa (rice) and the C4 plant Setaria viridis (green millet) revealed the roles of these genes in both minor vein formation and ground cell differentiation. Genetic and in vitro studies further suggested that SHR regulates this process through its interactions with IDD12 and 13. We also revealed direct interactions of these IDD proteins with a putative regulatory element within the auxin transporter gene PIN5c. Collectively, these findings indicate that a SHR–IDD regulatory circuit mediates auxin transport by negatively regulating PIN expression to modulate minor vein patterning in the grasses. A SHORT ROOT–INTERMEDIATE DOMAIN regulatory circuit modulates auxin transport by negatively regulating PIN-FORMED expression to regulate minor vein patterning in grasses. [ABSTRACT FROM AUTHOR]

Architectural plasticity in traits should be useful for understanding morphological and allocation plasticity at the whole-plant level and associated growth strategies of plants in dealing with variation in emergence time. To investigate how plants respond to emergence timing via dynamic architectural plasticity, we conducted a field experiment by sowing plants of Abutilon theophrasti on different dates to make them emerge in spring, late spring, summer and late summer, as four treatments of emergence time (ET1–ET4), and measured a series of mass and morphological traits in different vertical layers (with an inter-distance of 10 or 20 cm per layer) of plants at three stages. The 'vertical arch' and 'inverted-cone' shapes displayed at reproductive stage by plants emerged between spring and summer (ET1–ET3) and in late summer (ET4), respectively, revealed that they had contrasting timing schemes for vegetative and reproductive growth. Plants emerged earlier had relatively isolated vegetative and reproductive stages, while late-summer emerged plants had overlapped vegetative and reproductive stages due to accelerated leaf and reproductive growth simultaneously, as a result of shortened lifetime. In spite of this, plants emerged earlier were still able to adopt different optimal strategies in dealing with their corresponding circumstances. Architectural analysis should be a useful approach to better understanding diverse plant strategies in the context of varying environmental conditions. [ABSTRACT FROM AUTHOR]

Plants undergo extended morphogenesis. The shoot apical meristem (SAM) allows for reiterative development and the formation of new structures throughout the life of the plant. Intriguingly, the SAM produces morphologically different leaves in an age-dependent manner, a phenomenon known as heteroblasty. In Arabidopsis thaliana , the SAM produces small orbicular leaves in the juvenile phase, but gives rise to large elliptical leaves in the adult phase. Previous studies have established that a developmental decline of microRNA156 (miR156) is necessary and sufficient to trigger this leaf shape switch, although the underlying mechanism is poorly understood. Here we show that the gradual increase in miR156-targeted SQUAMOSA PROMOTER BINDING PROTEIN-LIKE transcription factors with age promotes cell growth anisotropy in the abaxial epidermis at the base of the leaf blade, evident by the formation of elongated giant cells. Time-lapse imaging and developmental genetics further revealed that the establishment of adult leaf shape is tightly associated with the longitudinal cell expansion of giant cells, accompanied by a prolonged cell proliferation phase in their vicinity. Our results thus provide a plausible cellular mechanism for heteroblasty in Arabidopsis, and contribute to our understanding of anisotropic growth in plants. [ABSTRACT FROM AUTHOR]

Leaves and flowers are crucial for the growth and development of higher plants. In this study we identified a mutant with narrow leaflets and early flowering (nlef) in an ethyl methanesulfonate-mutagenized population of woodland strawberry (Fragaria vesca) and aimed to identify the candidate gene. Genetic analysis revealed that a single recessive gene, nlef, controlled the mutant phenotype. We found that FvH4_1g25470, which encodes a putative DNA polymerase α with a polymerase and histidinol phosphatase domain (PHP), might be the candidate gene, using bulked segregant analysis with whole-genome sequencing, molecular markers, and cloning analyses. A splice donor site mutation (C to T) at the 5′ end of the second intron led to an erroneous splice event that reduced the expression level of the full-length transcript of FvePHP in mutant plants. FvePHP was localized in the nucleus and was highly expressed in leaves. Silencing of FvePHP using the virus-induced gene silencing method resulted in partial developmental defects in strawberry leaves. Overexpression of the FvePHP gene can largely restore the mutant phenotype. The expression levels of FveSEP1, FveSEP3, FveAP1, FveFUL, and FveFT were higher in the mutants than those in 'Yellow Wonder' plants, probably contributing to the early flowering phenotype in mutant plants. Our results indicate that mutation in FvePHP is associated with multiple developmental pathways. These results aid in understanding the role of DNA polymerase in strawberry development. [ABSTRACT FROM AUTHOR]

The response of plant leaf and root phenology and biomass in the Arctic to global change remains unclear due to the lack of synchronous measurements of above- and belowground parts. Our objective was to determine the phenological dynamics of the above- and belowground parts of Eriophorum vaginatum in the Arctic and its response to warming. We established a common garden located at Toolik Lake Field Station; tussocks of E. vaginatum from three locations, Coldfoot, Toolik Lake and Sagwon, were transplanted into the common garden. Control and warming treatments for E. vaginatum were set up at the Toolik Lake during the growing seasons of 2016 and 2017. Digital cameras, a handheld sensor and minirhizotrons were used to simultaneously observe leaf greenness, normalized difference vegetation index and root length dynamics, respectively. Leaf and root growth rates of E. vaginatum were asynchronous such that the timing of maximal leaf growth (mid-July) was about 28 days earlier than that of root growth. Warming of air temperature by 1 °C delayed the timing of leaf senescence and thus prolonged the growing season, but the temperature increase had no significant effect on root phenology. The seasonal dynamics of leaf biomass were affected by air temperature, whereas root biomass was correlated with soil thaw depth. Therefore, we suggest that leaf and root components should be considered comprehensively when using carbon and nutrient cycle models, as above- and belowground productivity and functional traits may have a different response to climate warming. [ABSTRACT FROM AUTHOR]

Hybrids between Arabidopsis thaliana accessions are important in revealing the consequences of epistatic interactions in plants. F1 hybrids between the A. thaliana accessions displaying either defense or developmental phenotypes have been revealing the roles of the underlying epistatic genes. The interaction of two naturally occurring alleles of the OUTGROWTH - ASSOCIATED KINASE (OAK) gene in Sha and Lag2-2, previously shown to cause a similar phenotype in a different allelic combination in A. thaliana , was required for the hybrid phenotype. Outgrowth formation in the hybrids was associated with reduced levels of salicylic acid, jasmonic acid and abscisic acid in petioles and the application of these hormones mitigated the formation of the outgrowths. Moreover, different abiotic stresses were found to mitigate the outgrowth phenotype. The involvement of stress and hormone signaling in outgrowth formation was supported by a global transcriptome analysis, which additionally revealed that TCP1 , a transcription factor known to regulate leaf growth and symmetry, was downregulated in the outgrowth tissue. These results demonstrate that a combination of natural alleles of OAK regulates growth and development through the integration of hormone and stress signals and highlight the importance of natural variation as a resource to discover the function of gene variants that are not present in the most studied accessions of A. thaliana. [ABSTRACT FROM AUTHOR]

Carbon and water are two main factors limiting leaf expansion. Restriction of leaf growth by low availability of carbon or water is among the earliest visible effects of potassium (K) deficiency. It is not known how K is involved in regulating the rhythmic supply of these two substrates, which differ remarkably across the day–night cycle, affecting leaf expansion. We investigated the effects of different K regimes on the time courses of leaf expansion, carbon assimilation, carbohydrates, and hydraulic properties of Brassica napus. Potassium supply increased leaf area, predominantly by promoting night-time leaf expansion (>60%), which was mainly associated with increased availability of carbohydrates from photosynthetic carbon fixation and import from old leaves rather than improvement of leaf hydraulics. However, sufficient K improved leaf hydraulic conductance to balance diurnal evaporative water loss and increase the osmotic contribution of water-soluble carbohydrates, thereby maintaining leaf turgor and increasing the daytime expansion rate. The results also indicated an ontogenetic role of K in modifying the amplitude of circadian expansion; almost 80% of the increase in leaf area occurred before the area reached 66.9% of the mature size. Our data provide mechanistic insight into K-mediated diel coordination of rhythmic carbon supply and water balance in leaf expansion. [ABSTRACT FROM AUTHOR]

Leaves, being a key plant organ involved in photosynthesis, play an important role in plant growth and development. Although there have been a few studies on the effects of potassium (K+) deficiency on the leaves of woody plants, knowledge about mechanism of necrotic spot formation on leaves during K+ deficiency is scarce. We used a hydroponics setup to understand the effects of K+ deficiency on Neolamarckia cadamba (Roxb.) Bosser. K+ deficiency resulted in smaller leaves and necrotic spots on the older leaves, whereas regulatory modules of the differentially expressed genes (DEGs) involved in cell proliferation, cell cycle and cell expansion were downregulated. K+ deficiency increased the activity of reactive oxygen species scavenging enzymes such as superoxide dismutase, ascorbate peroxidases and malondialdehyde, and expression of DEGs related to these was also upregulated. Strong diaminobenzidine staining was observed on the older leaves showing accumulation of H2O2 during K+ deficiency treatment. In addition, putrescine and ethylene synthesis genes were upregulated. Fifteen DEGs in response to ethylene signaling, including ETR1 , ETR2 , EBF1 , ERF1 and ERF2 , were upregulated in the third week. The leaf growth changes caused by K+ deficiency in N. cadamba were well demonstrated by our findings. [ABSTRACT FROM AUTHOR]

Phosphorus (P) is an essential nutrient for plants. Membrane lipid remodeling is an adaptive mechanism for P-starved plants that replaces membrane phospholipids with non-P galactolipids, presumably to retrieve scarce P sources and maintain membrane integrity. Whereas metabolic pathways to convert phospholipids to galactolipids are well-established, the mechanism by which phospholipid biosynthesis is involved in this process remains elusive. Here, we report that phospho-base N -methyltransferases 1 and 2 (PMT1 and PMT2), which convert phosphoethanolamine to phosphocholine (PCho), are transcriptionally induced by P starvation. Shoots of seedlings of pmt1 pmt2 double mutant showed defective growth upon P starvation; however, membrane lipid profiles were unaffected. We found that P-starved pmt1 pmt2 with defective leaf growth had reduced PCho content, and the growth defect was rescued by exogenous supplementation of PCho. We propose that PMT1 and PMT2 are induced by P starvation to produce PCho mainly for leaf growth maintenance, rather than for phosphatidylcholine biosynthesis, in membrane lipid remodeling. [ABSTRACT FROM AUTHOR]

Leaf laminar growth and adaxial–abaxial boundary formation are fundamental outcomes of plant development. Boundary and laminar growth coordinate the further patterning and growth of the leaf, directing the differentiation of cell types within the top and bottom domains and promoting initiation of lateral organs along their adaxial or abaxial axis. Leaf adaxial–abaxial polarity specification and laminar outgrowth are regulated by two transcription factors, REVOLUTA (REV) and KANADI (KAN). ABA INSENSITIVE TO GROWTH 1 (ABIG1) encodes a HOMEODOMAIN-LEUCINE ZIPPER (HD-ZIP) class II transcription factor and is a direct target of the adaxial–abaxial regulators REV and KAN. To investigate the role of ABIG1 in leaf development and in the establishment of polarity, we examined the phenotypes of both gain-of-function and loss-of-function mutants. Through genetic interaction analysis with REV and KAN mutants, we determined that ABIG1 plays a role in leaf laminar growth as well as in adaxial–abaxial polarity establishment. Genetic and physical interaction assays showed that ABIG1 interacts with the transcriptional TOPLESS corepressor. This study provides new evidence that ABIG1, another HD-ZIP II, facilitates growth through the corepressor TOPLESS. [ABSTRACT FROM AUTHOR]

The effect of nitrogen (N) nutrition on grapevine carbon (C) dynamics has been well studied at the annual scale, but poorly addressed at a pluriannual timescale. The aim of this study was to quantify, in an integrated conceptual framework, the effect of N nutrition on potted grapevine growth and storage over 2 consecutive years. The consequences of using destructive measurements were investigated using a hierarchical Bayesian model. The rate and duration of leaf growth were both positively impacted by the chlorophyll content of the leaves, but they were negatively impacted by the initial carbohydrate measurements, raising a distortion in the estimation of initial reserves. The C production per unit of global radiation depended on the leaf area dynamics. The allocation of dry matter mainly relied on the phenological stage. The present study highlights the importance of using appropriate statistical methods to overcome uncertainties due to destructive measurements. The genericity of the statistical approach presented may encourage its implementation in other agronomy studies. Based on our results, a simple conceptual framework of grapevine pluriannual growth under various N supplies was built. This provides a relevant basis for a future model of C and N balance and responses to N fertilization in grapevine. [ABSTRACT FROM AUTHOR]

Purine degradation products have been shown to play roles in plant response to stresses such as drought, salinity, extended dark, nitrogen deficiency, and pathogen infection. In this study, we used Arabidopsis wild-type (WT) and an Atxdh1 -knockout mutant defective in xanthine dehydrogenase1 (XDH1) to examine the role of degraded purine metabolites in the responses to wounding or UV-C stress applied to the middle leaves of the plant. Wounding or UV-C stress in the mutant resulted in lower fresh-weight, increased senescence symptoms, and increased cell death compared to WT plants. In addition, WT plants exhibited lower levels of oxidative stress indicators, reactive oxygen species, and malondialdehyde in their leaves than the mutant. Notably, transcripts and proteins functioning in the purine degradation pathway were regulated in such a way that it led to enhanced ureide levels in WT leaves 24h after applying the UV-C or wound stress. However, different remobilization of the accumulated ureides was observed after 72h of stress. In plants treated with UV-C, the concentration of allantoin was highest in young leaves, whereas in wounded plants it was lowest in these leaves and instead accumulated mainly in the middle leaves that had been wounded. These results indicated that in WT plants treated with UV-C, ureides were remobilized from the lower older and damaged leaves to support young leaf growth during the recovery period from stress. After wounding, however, whilst some ureides were remobilized to the young leaves, more remained in the wounded middle leaves to function as antioxidants and/or healing agents. [ABSTRACT FROM AUTHOR]

Background and Aims Mangrove plants are mostly found in tropical and sub-tropical tidal flats, and their limited distribution may be related to their responses to growth temperatures. However, the mechanisms underlying these responses have not been clarified. Here, we measured the dependencies of the growth parameters and respiration rates of leaves and roots on growth temperatures in typical mangrove species. Methods We grew two typical species of Indo-Pacific mangroves, Bruguiera gymnorrhiza and Rhizophora stylosa , at four different temperatures (15, 20, 25 and 30 °C) by irrigating with fresh water containing nutrients, and we measured growth parameters, chemical composition, and leaf and root O2 respiration rates. We then estimated the construction costs of leaves and roots and the respiration rates required for maintenance and growth. Key Results The relative growth rates of both species increased with growth temperature due to changes in physiological parameters such as net assimilation rate and respiration rate rather than to changes in structural parameters such as leaf area ratio. Both species required a threshold temperature for growth (12.2 °C in B. gymnorrhiza and 18.1 °C in R. stylosa). At the low growth temperature, root nitrogen uptake rate was lower in R. stylosa than in B. gymnorrhiza , leading to a slower growth rate in R. stylosa. This indicates that R. stylosa is more sensitive than B. gymnorrhiza to low temperature. Conclusions Our results suggest that the mangrove species require a certain warm temperature to ensure respiration rates sufficient for maintenance and growth, particularly in roots. The underground temperature probably limits their growth under the low-temperature condition. The lower sensitivity of B. gymnorrhiza to low temperature shows its potential to adapt to a wider habitat temperature range than R. stylosa. These growth and respiratory features may explain the distribution patterns of the two mangrove species. [ABSTRACT FROM AUTHOR]

There are potential sources of alleles and genes currently present in wheat-related species that have the potential to be introduced into wheat breeding programs targeting current and future hot and dry climates. However, to date neither the intra- nor the interspecific diversity of the responses of leaf growth and transpiration to temperature and evaporative demand have been investigated in across a significant range of wheat-related species. By analysing 12 groups of wheat-related species and subspecies, we were able to examine the multi-dimensional structure of the genetic diversity for traits linked to plant vegetative structures and their development, and to leaf expansion and transpiration, together with their responses to 'non-stressing' ranges of temperature and evaporative demand. In addition to providing new insights on how genome type, ploidy level, phylogeny, and breeding pressure act together to structure this genetic diversity, our study also provides new mathematical formalisms and associated parameters for trait responses across a wide range of genetic diversity in wheat-related species. This will potentially allow crop models to predict the impact of this diversity on yield, and thus to indicate potential sources of varietal improvement for modern wheat germplasms through interspecific crosses. [ABSTRACT FROM AUTHOR]

In nature, plants are usually subjected to a light/temperature regime of warm days and cold nights (referred to as +DIF). Compared with growth under +DIF, Arabidopsis plants show compact growth under the same photoperiod, but with an inverse temperature regime (cold days and warm nights: –DIF). Here we show that –DIF differentially affects the phase and amplitude of core clock gene expression. Under –DIF, the phase of the morning clock gene CIRCADIAN CLOCK ASSOCIATED 1 (CCA1) is delayed, similar to that of plants grown on low sucrose. Indeed, under –DIF, carbohydrate (CHO) starvation marker genes are specifically up-regulated at the end of the night (EN) in Arabidopsis rosettes. However, only in inner rosette tissue (small sink leaves and petioles of older leaves) are sucrose levels lower under –DIF compared with under +DIF, suggesting that sucrose in source leaf blades is not sensed for CHO status and that sucrose transport from source to sink may be impaired at EN. CHO starvation under –DIF correlated with increased starch breakdown during the night and decreased starch accumulation during the day. Moreover, we demonstrate that different ways of inducing CHO starvation all link to reduced growth of sink leaves. Practical implications for control of plant growth in horticulture are discussed. [ABSTRACT FROM AUTHOR]

Background and Aims Carbon reserves are a critical source of energy and substrates that allow trees to cope with periods of minimal carbon gain and/or high carbon demands, conditions which are prevalent in high-latitude forests. However, we have a poor understanding of carbon reserve dynamics at the whole-tree level in mature boreal trees. We therefore sought to quantify the seasonal changes in whole-tree and organ-level carbon reserve pools in mature boreal Betula papyrifera. Methods Non-structural carbohydrate (NSC; soluble sugars and starch) tissue concentrations were measured at key phenological stages throughout a calendar year in the roots, stem (inner bark and xylem), branches and leaves, and scaled up to estimate changes in organ and whole-tree NSC pool sizes. Fine root and stem growth were also measured to compare the timing of growth processes with changes in NSC pools. Key Results The whole-tree NSC pool increased from its spring minimum to its maximum at bud set, producing an average seasonal fluctuation of 0.96 kg per tree. This fluctuation represents a 72 % change in the whole-tree NSC pool, which greatly exceeds the relative change reported for more temperate conspecifics. At the organ level, branches accounted for roughly 48–60 % of the whole-tree NSC pool throughout the year, and their seasonal fluctuation was four to eight times greater than that observed in the stemwood, coarse roots and inner bark. Conclusions Branches in boreal B. papyrifera were the largest and most dynamic storage pool, suggesting that storage changes at the branch level largely drive whole-tree storage dynamics in these trees. The greater whole-tree seasonal NSC fluctuation in boreal vs. temperate B. papyrifera may result from (1) higher soluble sugar concentration requirements in branches for frost protection, and/or (2) a larger reliance on reserves to fuel new leaf and shoot growth in the spring. [ABSTRACT FROM AUTHOR]

Plant cell deformations are driven by cell pressurization and mechanical constraints imposed by the nanoscale architecture of the cell wall, but how these factors are controlled at the genetic and molecular levels to achieve different types of cell deformation is unclear. Here, we used stomatal guard cells to investigate the influences of wall mechanics and turgor pressure on cell deformation and demonstrate that the expression of the pectin-modifying gene PECTATE LYASE LIKE12 (PLL12) is required for normal stomatal dynamics in Arabidopsis thaliana. Using nanoindentation and finite element modeling to simultaneously measure wall modulus and turgor pressure, we found that both values undergo dynamic changes during induced stomatal opening and closure. PLL12 is required for guard cells to maintain normal wall modulus and turgor pressure during stomatal responses to light and to tune the levels of calcium crosslinked pectin in guard cell walls. Guard cell-specific knockdown of PLL12 caused defects in stomatal responses and reduced leaf growth, which were associated with lower cell proliferation but normal cell expansion. Together, these results force us to revise our view of how wall-modifying genes modulate wall mechanics and cell pressurization to accomplish the dynamic cellular deformations that underlie stomatal function and tissue growth in plants. [ABSTRACT FROM AUTHOR]

DORMANCY-ASSOCIATED MADS-box (DAM) and SHORT VEGETATIVE PHASE (SVP) genes have been implicated in the regulation of winter dormancy in perennials. Ectopic expression of apple (Malus × domestica Borkh. 'Royal Gala') DAM and SVP genes delays budbreak and constrains lateral shoot outgrowth. In this study, we used RNA interference (RNAi) to simultaneously target all apple DAM and SVP genes in order to study their role and mode of action in the regulation of bud dormancy, budbreak and flowering. A synthetic construct carrying a hairpin fragment assembled from sequences specific to coding regions of three DAM and two SVP genes was used to generate transgenic lines. Reduced expression of DAM/SVP genes resulted in delayed leaf senescence and abscission in autumn, failure to enter bud dormancy in winter and continual growth of new leaves regardless of the season for over 3 years. Precocious flowering but normal flower morphology, fertility and fruit development were observed. The non-dormant phenotype was associated with modified phytohormone composition. The content of gibberellins (GAs) and jasmonates (JAs) was significantly increased in terminal buds of RNAi lines compared with wildtype plants, accompanied by elevated expression of the key GA biosynthesis pathway gene GIBBERELLIN 20 OXIDASE-2 (MdGA20ox-2) along with the FLOWERING LOCUS T gene MdFT2. The key mediator of plasmodesmatal closure, MdCALLOSE SYNTHASE 1 (MdCALS1), was repressed in RNAi lines. This study provides functional evidence for the role of DAM/SVP genes in vegetative phenology of apple and paves the way for production of low-chill varieties suitable for growth in warming climates. [ABSTRACT FROM AUTHOR]

Salt stress is considered one of the most devastating environmental stresses, affecting barley growth and leading to significant yield loss. Hence, there is considerable interest in investigating the most effective traits that determine barley growth under salt stress. The objective of this study was to elucidate the contribution of osmotic and oxidative stress components in leaves and roots growth under salt stress. Two distinct barley (Hordeum vulgare) salt-stress tolerant genotypes, Barrage Malleg (BM, tolerant) and Saouef (Sf, sensitive), were subjected to 200 mM NaCl at early vegetative stages. Stressed and control leaves and roots tissue were assessed for several growth traits, including fresh and dry weight and plant length, as well as the content of osmoprotectants proline and soluble sugars. In addition, malondialdehyde content and activities of superoxide dismutase (SOD), catalase (CAT) and ascorbate peroxidase (APX), as well as their corresponding gene expression patterns, were investigated. The results showed better performance of BM over Sf for leaf dry weight (LDW), root dry weight (RDW) and root length (RL). The salt-tolerant genotype (BM) had better osmoprotection against salt stress compared with the salt-sensitive genotype (Sf), with a higher accumulation of proline and soluble sugars in leaves and roots and a stronger antioxidant system as evidenced by higher activities of SOD, CAT and APX and more abundant Cu/Zn-SOD transcripts, especially in roots. Stepwise regression analysis indicated that under salt stress the most predominant trait of barley growth was Cu/Zn-SOD gene expression level, suggesting that alleviating oxidative stress and providing cell homeostasis is the first priority. [ABSTRACT FROM AUTHOR]

Reduced blue light irradiance is known to enhance leaf elongation rate (LER) in grasses, but the mechanisms involved have not yet been elucidated. We investigated whether leaf elongation response to reduced blue light could be mediated by stomata-induced variations of plant transpiration. Two experiments were carried out on tall fescue in order to monitor LER and transpiration under reduced blue light irradiance. Additionally, LER dynamics were compared with those observed in the response to vapour pressure deficit (VPD)-induced variations of transpiration. Finally, we developed a model of water flow within a tiller to simulate the observed short-term response of LER to various transpiration regimes. LER dramatically increased in response to blue light reduction and then reached new steady states, which remained higher than the control. Reduced blue light triggered a simultaneous stomatal closure which induced an immediate decrease of leaf transpiration. The hydraulic model of leaf elongation accurately predicted the LER response to blue light and VPD, resulting from an increase in the growth-induced water potential gradient in the leaf growth zone. Our results suggest that the blue light signal is sensed by stomata of expanded leaves and transduced to the leaf growth zone through the hydraulic architecture of the tiller. [ABSTRACT FROM AUTHOR]

Several breeding initiatives have sought to improve flag leaf performance as its health and physiology are closely correlated to rice yield. Previous studies have described natural variation of photosynthesis for flag leaves; however, none has examined their performance under the non-steady-state conditions that prevail in crop fields. Photosynthetic induction is the transient response of photosynthesis to a change from low to high light. Rice flag leaf photosynthesis was measured in both steady- and non-steady-state conditions to characterize natural variation. Between the lowest and highest performing accession, there was a 152% difference for average CO2 assimilation during induction (Ā 300), a 77% difference for average intrinsic water use efficiency during induction (iWUEavg), and a 185% difference for the speed of induction (IT50), indicating plentiful variation. No significant correlation was found between steady- and non-steady-state photosynthetic traits. Additionally, measures of neither steady-state nor non-steady-state photosynthesis of flag leaves correlated with the same measures of leaves in the vegetative growth stage, with the exception of iWUEavg. Photosynthetic induction was measured at six [CO2], to determine biochemical and diffusive limitations to photosynthesis in vivo. Photosynthetic induction in rice flag leaves was limited primarily by biochemistry. [ABSTRACT FROM AUTHOR]

Aims In plant eco-physiology, less negative (enriched) carbon 13 (13C) in the leaves indicates conditions of reducing leaf gas exchange through stomata, e.g. under drought. In addition, 13C is expected to be less negative in non-photosynthetic tissues as compared with leaves. However, these relationships in δ 13C from leaves (photosynthetic organs) to branches, stems and roots (non-photosynthetic organs) are rarely tested across multiple closely related tree species, multiple compartments, or in trees growing under extreme heat and drought. Methods We measured leaf-to-root 13C in three closely related desert acacia species (Acacia tortilis , A. raddiana and A. pachyceras). We measured δ 13C in leaf tissues from mature trees in southern Israel. In parallel, a 7-year irrigation experiment with 0.5, 1.0 or 4.0 L day−1 was conducted in an experimental orchard. At the end of the experiment, growth parameters and δ 13C were measured in leaves, branches, stems and roots. Important Findings The δ 13C in leaf tissues sampled from mature trees was ca. −27‰, far more depleted than expected from a desert tree growing in one of the Earth's driest and hottest environments. Across acacia species and compartments, δ 13C was not enriched at all irrigation levels (−28‰ to ca. −27‰), confirming our measurements in the mature trees. Among compartments, leaf δ 13C was unexpectedly similar to branch and root δ 13C, and surprisingly, even less negative than stem δ 13C. The highly depleted leaf δ 13C suggests that these trees have high stomatal gas exchange, despite growing in extremely dry habitats. The lack of δ 13C enrichment in non-photosynthetic tissues might be related to the seasonal coupling of growth of leaves and heterotrophic tissues. [ABSTRACT FROM AUTHOR]

BSD (mammalian B TF2-like transcription factors, s ynapse-associated proteins, and D OS2-like proteins) is a conserved domain that exists in a variety of organisms, but its function has not been well studied. Here, we identified a novel BSD domain-containing protein (SlBSD1) in tomato (Solanum lycopersicum). Biochemical and microscopy assays indicated that SlBSD1 is a functional transcription factor that is predominantly localized in the nucleus. Loss-of-function and overexpression analyses suggested that SlBSD1 is a novel regulator of vegetative growth and leaf senescence in tomato. SlBSD1 -knockdown (-KD) plants exhibited retarded vegetative growth and precocious leaf senescence, whereas SlBSD1 -overexpression (-OX) plants displayed the opposite phenotypes. The negative role of SlBSD1 in leaf senescence was also supported by RNA-seq analysis comparing leaf tissues from SlBSD1 -KD and wild-type plants. In addition, contents of soluble solids were altered in fruits in the SlBSD1 -KD and SlBSD1 -OX plants. Taken together, our data suggest that the novel transcription factor SlBSD1 plays important roles in controlling fruit quality and other physiological processes in tomato, including vegetative growth and leaf senescence. [ABSTRACT FROM AUTHOR]

The quality of yield prediction is linked to that of leaf area. We first analysed the consequences of flowering time and environmental conditions on the area of individual leaves in 127 genotypes presenting contrasting flowering times in fields of Europe, Mexico, and Kenya. Flowering time was the strongest determinant of leaf area. Combined with a detailed field experiment, this experiment showed a large effect of flowering time on the final leaf number and on the distribution of leaf growth rate and growth duration along leaf ranks, in terms of both length and width. Equations with a limited number of genetic parameters predicted the beginning, end, and maximum growth rate (length and width) for each leaf rank. The genotype-specific environmental effects were analysed with datasets in phenotyping platforms that assessed the effects (i) of the amount of intercepted light on leaf width, and (ii) of temperature, evaporative demand, and soil water potential on leaf elongation rate. The resulting model was successfully tested for 31 hybrids in 15 European and Mexican fields. It potentially allows prediction of the vertical distribution of leaf area of a large number of genotypes in contrasting field conditions, based on phenomics and on sensor networks. [ABSTRACT FROM AUTHOR]

Shoot architecture is a key component of the interactions between plants and their environment. We present a novel model of grass, which fully integrates shoot morphogenesis and the metabolism of carbon (C) and nitrogen (N) at organ scale, within a three-dimensional representation of plant architecture. Plant morphogenesis is seen as a self-regulated system driven by two main mechanisms. First, the rate of organ extension and the establishment of architectural traits are regulated by concentrations of C and N metabolites in the growth zones and the temperature. Second, the timing of extension is regulated by rules coordinating successive phytomers instead of a thermal time schedule. Local concentrations are calculated from a model of C and N metabolism at organ scale. The three-dimensional representation allows the accurate calculation of light and temperature distribution within the architecture. The model was calibrated for wheat (Triticum aestivum) and evaluated for early vegetative stages. This approach allowed the simulation of realistic patterns of leaf dimensions, extension dynamics, and organ mass and composition. The model simulated, as emergent properties, plant and agronomic traits. Metabolic activities of growing leaves were investigated in relation to whole-plant functioning and environmental conditions. The current model is an important step towards a better understanding of the plasticity of plant phenotype in different environments. [ABSTRACT FROM AUTHOR]

Dendrometers are being increasingly used to measure stem radius changes in trees and to unravel the mechanisms underlying stem daily rhythms of radial expansion and contraction. Nevertheless, automated dendrometers have not been often used to measure root radius dynamics, their relationship with environmental variables and the influence of endogenous processes, especially in drought-prone Mediterranean areas. Here, we measured root radius dynamics of two coexisting oak species (the evergreen Quercus ilex L. and the deciduous Quercus faginea Lam). Our goals were to describe annual, seasonal and diurnal scale root radius patterns and to disentangle the role of different environmental parameters as drivers. Long-term high-resolution measurements (every 15 min over 7 years) were collected with automated point dendrometers on the main tree roots of five individuals per species. Root radius annual change patterns were bimodal and similar for both oak species. Quercus faginea Lam showed three times larger root increment in the spring than Q. ilex , but the bimodal pattern was stronger in Q. ilex , which showed a larger root increment in autumn. Quercus faginea Lam showed an earlier root phenological activation in the spring and in late summer compared with Q. ilex. The effects of environmental drivers across species were similar at daily scales: root radius increased with air temperature and soil moisture, and it decreased with rising vapor pressure deficit. Furthermore, daily root radius variations for both oak species were maintained after extracting statistically the environmental effects, which points toward a significant role of endogenous drivers. These differences in root radius change patterns at seasonal to daily scales likely result from the differences in leaf phenology and growth strategy. Quercus faginea Lam is deciduous and has a faster growing rate in spring than the evergreen Q. ilex, which can grow more in summer. [ABSTRACT FROM AUTHOR]

An increased frequency of fire events on the Slovenian Karst is in line with future climate change scenarios for drought-prone environments worldwide. It is therefore of the utmost importance to better understand tree–fire–climate interactions for predicting the impact of changing environment on tree functioning. To this purpose, we studied the post-fire effects on leaf development, leaf carbon isotope composition (δ 13C), radial growth patterns and the xylem and phloem anatomy in undamaged (H-trees) and fire-damaged trees (F-trees) of Quercus pubescens Willd. with good resprouting ability in spring 2017, the growing season after a rangeland fire in August 2016. We found that the fully developed canopy of F-trees reached only half of the leaf area index values measured in H-trees. Throughout the season, F-trees were characterized by higher water potential and stomatal conductivity and achieved higher photosynthetic rates compared to unburnt H-trees. The foliage of F-trees had more negative δ 13C values than those of H-trees. This reflects that F-trees less frequently meet stomatal limitations due to reduced transpirational area and more favourable leaf-to-root ratio. In addition, the growth of leaves in F-trees relied more on the recent photosynthates than on reserves due to the fire disturbed starch accumulation in the previous season. Cambial production stopped 3 weeks later in F-trees, resulting in 60 and 22% wider xylem and phloem increments, respectively. A novel approach by including phloem anatomy in the analyses revealed that fire caused changes in conduit dimensions in the early phloem but not in the earlywood. However, premature formation of the tyloses in the earlywood vessels of the youngest two xylem increments in F-trees implies that xylem hydraulic integrity was also affected by heat. Analyses of secondary tissues showed that although xylem and phloem tissues are interlinked changes in their transport systems due to heat damage are not necessarily coordinated. [ABSTRACT FROM AUTHOR]

Due to its subtropical origins, rice (Oryza sativa) is sensitive to low-temperature stress. In this study, we identify LOC_Os04g24110, annotated to encode the UDP-glycosyltransferase enzyme UGT90A1, as a gene associated with the low-temperature seedling survivability (LTSS) quantitative trait locus qLTSS4-1. Differences between haplotypes in the control region of OsUGT90A1 correlate with chilling tolerance phenotypes, and reflect differential expression between tolerant and sensitive accessions rather than differences in protein sequences. Expression of OsUGT90A1 is initially enhanced by low temperature, and its overexpression helps to maintain membrane integrity during cold stress and promotes leaf growth during stress recovery, which are correlated with reduced levels of reactive oxygen species due to increased activities of antioxidant enzymes. In addition, overexpression of OsUGT90A1 in Arabidopsis improves freezing survival and tolerance to salt stress, again correlated with enhanced activities of antioxidant enzymes. Overexpression of OsUGT90A1 in rice decreases root lengths in 3-week-old seedlings while gene-knockout increases the length, indicating that its differential expression may affect phytohormone activities. We conclude that higher OsUGT90A1 expression in chilling-tolerant accessions helps to maintain cell membrane integrity as an abiotic stress-tolerance mechanism that prepares plants for the resumption of growth and development during subsequent stress recovery. [ABSTRACT FROM AUTHOR]

By modifying two genes involved in lipid biosynthesis and storage [cysteine oleosin (cys-OLE)/diacylglycerol O -acyltransferase (DGAT)], the accumulation of stable lipid droplets in perennial ryegrass (Lolium perenne) leaves was achieved. Growth, biomass allocation, leaf structure, gas exchange parameters, fatty acids, and water-soluble carbohydrates were quantified for a high-expressing cys-OLE/DGAT ryegrass transformant (HL) and a wild-type (WT) control grown under controlled conditions with 1–10 mM nitrogen (N) supply at ambient and elevated atmospheric CO2. A dramatic shift in leaf carbon (C) storage occurred in HL leaves, away from readily mobilizable carbohydrates and towards stable lipid droplets. HL exhibited an increased growth rate, mainly in non-photosynthetic organs, leading to a decreased leaf mass fraction. HL leaves, however, displayed an increased specific leaf area and photosynthetic rate per unit leaf area, delivering greater overall C capture and leaf growth at high N supply. HL also exhibited a greater photosynthesis response to elevated atmospheric CO2. We speculate that by behaving as uniquely stable microsinks for C, cys-OLE-encapsulated lipid droplets can reduce feedback inhibition of photosynthesis and drive greater C capture. Manipulation of many genes and gene combinations has been used to increase non-seed lipid content. However, the cys-OLE/DGAT technology remains the only reported case that increases plant biomass. We contrast cys-OLE/DGAT with other lipid accumulation strategies and discuss the implications of introducing lipid sinks into non-seed organs for plant energy homeostasis and growth. [ABSTRACT FROM AUTHOR]

By modifying two genes involved in lipid biosynthesis and storage [cysteine oleosin (cys-OLE)/diacylglycerol O -acyltransferase (DGAT)], the accumulation of stable lipid droplets in perennial ryegrass (Lolium perenne) leaves was achieved. Growth, biomass allocation, leaf structure, gas exchange parameters, fatty acids, and water-soluble carbohydrates were quantified for a high-expressing cys-OLE/DGAT ryegrass transformant (HL) and a wild-type (WT) control grown under controlled conditions with 1–10 mM nitrogen (N) supply at ambient and elevated atmospheric CO2. A dramatic shift in leaf carbon (C) storage occurred in HL leaves, away from readily mobilizable carbohydrates and towards stable lipid droplets. HL exhibited an increased growth rate, mainly in non-photosynthetic organs, leading to a decreased leaf mass fraction. HL leaves, however, displayed an increased specific leaf area and photosynthetic rate per unit leaf area, delivering greater overall C capture and leaf growth at high N supply. HL also exhibited a greater photosynthesis response to elevated atmospheric CO2. We speculate that by behaving as uniquely stable microsinks for C, cys-OLE-encapsulated lipid droplets can reduce feedback inhibition of photosynthesis and drive greater C capture. Manipulation of many genes and gene combinations has been used to increase non-seed lipid content. However, the cys-OLE/DGAT technology remains the only reported case that increases plant biomass. We contrast cys-OLE/DGAT with other lipid accumulation strategies and discuss the implications of introducing lipid sinks into non-seed organs for plant energy homeostasis and growth. [ABSTRACT FROM AUTHOR]

We analysed the cellular and molecular changes in the leaf growth zone of tolerant and sensitive rice varieties in response to suboptimal temperatures. Cold reduced the final leaf length by 35% and 51% in tolerant and sensitive varieties, respectively. Tolerant lines exhibited a smaller reduction of the leaf elongation rate and greater compensation by an increased duration of leaf growth. Kinematic analysis showed that cold reduced cell production in the meristem and the expansion rate in the elongation zone, but the latter was compensated for by a doubling of the duration of cell expansion. We performed iTRAQ proteome analysis on proliferating and expanding parts of the leaf growth zone. We identified 559 and 542 proteins, of which 163 and 210 were differentially expressed between zones, and 96 and 68 between treatments, in the tolerant and sensitive lines, respectively. The categories protein biosynthesis and redox homeostasis were significantly overrepresented in the up-regulated proteins. We therefore measured redox metabolites and enzyme activities in the leaf growth zone, demonstrating that tolerance of rice lines to suboptimal temperatures correlates with the ability to up-regulate enzymatic antioxidants in the meristem and non-enzymatic antioxidants in the elongation zone. [ABSTRACT FROM AUTHOR]

Climate change will increase the risk of flooding in several areas of the world where Populus deltoides Marshall (eastern cottonwood) is planted, so it would be desirable for this species to select for flooding tolerance. The aims of this work were to explore the variability in growth, leaf traits and flooding tolerance in an F1 full-sib intraspecific progeny of P. deltoides , to analyze the correlations of leaf and growth traits with flooding tolerance and to assess their suitability for use in breeding programs. Two-month-old parental clones and their progeny of 30 full-sib F1 genotypes were grown in pots and subjected to two treatments: (i) plants watered to field capacity (control) and (ii) plants flooded up to 10 cm above soil level for 35 days. Growth (height, diameter and biomass partition) and leaf traits (leaf size and number, specific leaf area, leaf senescence, abscission, stomatal conductance, carbon isotope discrimination, stomatal index) were measured. Flooding tolerance for each genotype was estimated as the ratio of the biomass of stressed plants to the biomass of control plants. Results showed segregation in terms of flooding tolerance in the F1 progeny. A significant genotype effect was found for leaf size and number, carbon isotopic discrimination and stomatal conductance, but it did not correlate with flooding tolerance. Height, diameter and root-to-shoot ratio had a positive phenotypic correlation with flooding tolerance, and there was a positive genetic correlation of height and diameter with biomass on both treatments. The narrow sense heritability values for the traits analyzed ranged from 0 to 0.56. We conclude that growth traits are more adequate than leaf traits for selection to increase flooding tolerance. A vigorous initial growth would increase flooding tolerance in young poplar plants. [ABSTRACT FROM AUTHOR]

Background and aims Phenology and morphology are two major aspects of crop growth models. A new process-based model built for hardneck garlic (Allium sativum) is presented, focusing on phenology and morphology processes and how they translate to whole-plant growth. The tight coupling between the two processes and their dynamic changes throughout the growing season were captured while incorporating storage effects and reproductive aspects unique to bulbous crops. Methods Non-linear temperature dependences of leaf development were integrated into the model and dynamically coupled with changes in leaf growth throughout the growing season. Bulb storage effects on leaf development and photoperiod effects on the vegetative-to-reproductive transition were also incorporated. The model was parameterized with data from a set of experiments and the literature, while its performance was tested with additional observations that had not been used for parameterization under a range of environmental conditions, management practices and cultivar choices. Key Results The model successfully captured the dynamic nature of leaf development and growth in garlic plants throughout the growing season. It simulated with reasonable accuracy the timing of leaf initiation, maturation and senescence, as well as changes in green leaf area over time. Most parameters were relatively stable across cultivars, and parameter sensitivity tests revealed the importance of bulb storage effects. Conclusions The model embodies a novel approach to capture the phenology and morphology of garlic under a range of environments, genotypes and management practices. The process-oriented nature of the model and inclusion of storage effects set the foundation for bulbous crop growth simulations, allowing the understanding and discovery of key processes that coordinate and integrate the dynamics of growth and development from organ to whole plant, with implications for crop improvement programmes while opening opportunities for modelling other bulbous crops. [ABSTRACT FROM AUTHOR]

In this study, we investigated the regulatory function of miR396 in the phase transition in Arabidopsis thaliana. Using AtMIR396a/b knockout mutants generated through clustered regularly interspaced short palindromic repeats/CRISPR-associated protein 9 (CRISPR/Cas9)-directed genome editing, we showed that miR396 negatively regulates the leaf size and vegetative phase transition, and the first leaf with abaxial trichomes appeared earlier in the mir396ab double mutant than in the wild type (WT) and was significantly delayed in miR396 overexpression lines. Moreover, mir396ab exhibited early flowering, whereas 35S:MIR396a/b and cib4-1 delayed flowering, and the flowering time was negatively correlated with FT gene expression. Furthermore, in arp6 and pie1 mutants, which are deficient in the ATP-dependent chromatin remodeling complex (SWR1-C), miR396 expression was significantly repressed. Compared with the WT, reduced H2A.Z deposit and stronger relative nucleosome occupancy in the promoter region of MIR396a was found in the arp6 mutant, indicating that SWR1-C contributes to the transcriptional activation of MIR396a via nucleosome dynamics. In addition, miR396 displayed specific spatio-temporal expression patterns in the leaf, which was altered in arp6 and pie1 , and therefore affected the transcript levels of CIB4 and FT in these mutants. We propose that miR396 is not only a marker of cell differentiation, but also an age signal for leaf development and phase change. Meanwhile, SWR1-C-mediated epigenetic regulation contributes to the age-dependent enhancement of miR396 expression and differential miR396 accumulation among leaves. [ABSTRACT FROM AUTHOR]

Background and Aims The development of Arundo donax as a biomass crop for use on drought-prone marginal lands in areas with warm to hot climates is constrained by the lack of variation within this species. We investigated the effect of morphological and physiological variation on growth and tolerance to drought under field conditions in three ecotypes of A. donax collected from habitats representing a climate gradient: a pre-desert in Morocco, a semi-arid Mediterranean climate in southern Italy and a warm sub-humid region of central Italy. Methods The three A. donax ecotypes were grown under irrigated and rain-fed conditions in a common garden field trial in a region with a semi-arid Mediterranean climate. Physiological and morphological characteristics, and carbohydrate metabolism of the ecotypes were recorded to establish which traits were associated with yield and/or drought tolerance. Key Results Variation was observed between the A. donax ecotypes. The ecotype from the most arid habitat produced the highest biomass yield. Stem height and the retention of photosynthetic capacity later in the year were key traits associated with differences in biomass yield. The downregulation of photosynthetic capacity was not associated with changes in foliar concentrations of sugars or starch. Rain-fed plants maintained photosynthesis and growth later in the year compared with irrigated plants that began to senescence earlier, thus minimizing the difference in yield. Effective stomatal control prevented excessive water loss, and the emission of isoprene stabilized photosynthetic membranes under drought and heat stress in A. donax plants grown under rain-fed conditions without supplementary irrigation. Conclusions Arundo donax is well adapted to cultivation in drought-prone areas with warm to hot climates. None of the A. donax ecotypes exhibited all of the desired traits consistent with an 'ideotype'. Breeding or genetic (identification of quantitative trait loci) improvement of A. donax should select ecotypes on the basis of stem morphology and the retention of photosynthetic capacity. [ABSTRACT FROM AUTHOR]

Aims Anthocyanin accumulation is the main factor underlying why young plant leaves turn red, and plant growth follows the principle of maximizing the economic efficiency of energy. There is a need to verify the role of young plant leaves turning red and confirm whether anthocyanin accumulation overconsumes the energy of the plant. Methods We compared the different pigment contents, antioxidant capacities, leaf mass per area, photosynthetic characteristics, dark respiration rates, light compensation points (LCPs) and electron flow distribution of young and mature Schima superba leaves grown under full sunlight and 30% full sunlight conditions. We then examined the correlations between anthocyanins and total antioxidant capacity, photosynthetic pigments, dark respiration rates, LCPs by using linear regression. Finally, we analyzed Pearson correlation coefficients and used principal component analysis to evaluate the interactions of these functional indicators. Important Findings The young leaves of S. superba accumulated anthocyanins in full sunlight but not in 30% full sunlight. Anthocyanins substantially contributed to the total antioxidant capacity (accounting for 33.29%) in juvenile S. superba leaves. Young leaves containing more anthocyanin accumulated less reactive oxygen species after high light (HL) treatment. However, juvenile leaves accumulating anthocyanins showed 56.97% higher dark respiration rates, 24.79% higher LCPs and 5.32% higher allocations of electron flow to photorespiration and lower organic substance accumulation than did those without anthocyanins. These results suggest that young S. superba leaves must consume more energy to accumulate anthocyanins to adapt to HL environments. Therefore, plants sacrifice growth rate to ensure survival, which is a trade-off for their ability to respond to external environments. [ABSTRACT FROM AUTHOR]