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The polar auxin transport is required for proper plant growth and development. D6 PROTEIN KINASE (D6PK) is required for the phosphorylation of PIN‐FORMED (PIN) auxin efflux carriers to regulate auxin transport, while the regulation of D6PK stabilization is still poorly understood. Here, we found that Cytosolic ABA Receptor Kinases (CARKs) redundantly interact with D6PK, and the interactions are dependent on CARKs' kinase activities. Similarly, CARK3 also could interact with paralogs of D6PK, including D6PKL1, D6PKL2, and D6PKL3. The genetic analysis shows that D6PK acts the downstream of CARKs to regulate Arabidopsis growth, including hypocotyl, leaf area, vein formation, and the length of silique. Loss‐of‐function of CARK3 in overexpressing GFP‐D6PK plants leads to reduce the level of D6PK protein, thereby rescues plant growth. In addition, the cell‐free degradation assays indicate that D6PK is degraded through 26 S proteasome pathway, while the phosphorylation by CARK3 represses this process in cells. In summary, D6PK stabilization by the CARK family is required for auxin‐mediated plant growth and development. Summary statement: The AGC VIII kinase D6 PROTEIN KINASE (D6PK) phosphorylates PIN‐FORMED (PIN) auxin efflux carriers, which is essential for auxin transport in plant cells. This study uncovers a new mechanism for D6PK stability through Cytosolic ABA Receptor Kinases‐mediated phosphorylation to regulate auxin‐controlled Arabidopsis growth and development. [ABSTRACT FROM AUTHOR]

Plants contain three NADPH‐thioredoxin reductases (NTR) located in the cytosol/mitochondria (NTRA/B) and the plastid (NTRC) with important metabolic functions. However, mutants deficient in all NTRs remained to be investigated. Here, we generated and characterised the triple Arabidopsis ntrabc mutant alongside with ntrc single and ntrab double mutants under different environmental conditions. Both ntrc and ntrabc mutants showed reduced growth and substantial metabolic alterations, especially in sink leaves and under high CO2 (HC), as compared to the wild type. However, ntrabc showed higher effective quantum yield of PSII under both constant and fluctuating light conditions, altered redox states of NADH/NAD+ and glutathione (GSH/GSSG) and lower potential quantum yield of PSII in sink leaves in ambient but not high CO2 concentrations, as compared to ntrc, suggesting a functional interaction between chloroplastic and extra‐chloroplastic NTRs in photosynthesis regulation depending on leaf development and environmental conditions. Our results unveil a previously unknown role of the NTR system in regulating sink leaf metabolism and plant acclimation to HC, while it is not affecting full plant development, indicating that the lack of the NTR system can be compensated, at least to some extent, by other redox mechanisms. Summary Statement: We unveil that NTRC is crucial for sink leaf metabolism and plant acclimation to high CO2. Although Arabidopsis plants lacking all NADPH‐thioredoxin reductases (NTRs) have severely reduced growth, they produced viable seeds, indicating that NTRs are not essential for plant development and compensated by other redox mechanisms. [ABSTRACT FROM AUTHOR]

Maize (Zea mays), a cold‐sensitive crop, requires cold tolerance for extending the length of the growing season in temperate climates. However, response curves to different cold temperatures and exposure durations are lacking. We used a meta‐analysis approach using data from literature to investigate the effect of cold stress in the maize leaf. We constructed response curves to temperature and exposure durations for 18 key parameters related to leaf growth, photosynthesis, oxidative stress, antioxidants, and the phytohormone ABA. To determine their relevance for cold tolerance, we compared cold tolerant Flint and cold sensitive Dent lines. Treatment temperatures ranged from −20°C to 20°C for cold and from 12°C to 30°C for control and exposure duration from 3 min to 60 days. We found interacting effects of temperature and exposure durations on different response parameters. The strongest difference between Flint and Dent was observed for electrolyte leakage (EL). Our results show that the commonly used 4°C for cold and 25°C for control with medium cold exposure (1−7 days) induces a 50% decrease in shoot dry weight and leaf area and that EL is an easy and reliable indicator for cold tolerance studies. Summary Statement: Meta‐analysis of cold stress responses in the Zea mays leaf reveals differences in response curves of physiological parameters and shows that electrolyte leakage is a reliable indicator for cold tolerance. [ABSTRACT FROM AUTHOR]

Plants contain three NADPH‐thioredoxin reductases (NTR) located in the cytosol/mitochondria (NTRA/B) and the plastid (NTRC) with important metabolic functions. However, mutants deficient in all NTRs remained to be investigated. Here, we generated and characterised the triple Arabidopsis ntrabc mutant alongside with ntrc single and ntrab double mutants under different environmental conditions. Both ntrc and ntrabc mutants showed reduced growth and substantial metabolic alterations, especially in sink leaves and under high CO2 (HC), as compared to the wild type. However, ntrabc showed higher effective quantum yield of PSII under both constant and fluctuating light conditions, altered redox states of NADH/NAD+ and glutathione (GSH/GSSG) and lower potential quantum yield of PSII in sink leaves in ambient but not high CO2 concentrations, as compared to ntrc, suggesting a functional interaction between chloroplastic and extra‐chloroplastic NTRs in photosynthesis regulation depending on leaf development and environmental conditions. Our results unveil a previously unknown role of the NTR system in regulating sink leaf metabolism and plant acclimation to HC, while it is not affecting full plant development, indicating that the lack of the NTR system can be compensated, at least to some extent, by other redox mechanisms. Summary Statement: We unveil that NTRC is crucial for sink leaf metabolism and plant acclimation to high CO2. Although Arabidopsis plants lacking all NADPH‐thioredoxin reductases (NTRs) have severely reduced growth, they produced viable seeds, indicating that NTRs are not essential for plant development and compensated by other redox mechanisms. [ABSTRACT FROM AUTHOR]

Plants adapt to high temperature stresses through thermomorphogenesis, a process that includes stem elongation and hyponastic leaf growth. Thermomorphogenesis is gated by the circadian clock through two evening‐expressed clock components, TIMING OF CAB EXPRESSION1 (TOC1) and PSEUDO‐RESPONSE REGULATORS5 (PRR5). These proteins directly interact with and inhibit PHYTOCHROME INTERACTING FACTOR4 (PIF4), a basic helix–loop–helix transcription factor that promotes thermoresponsive growth. PIF4‐mediated thermoresponsive growth is positively regulated by ZEITLUPE (ZTL), a central clock component, but the molecular mechanisms underlying this are poorly understood. Here, we show that ZTL regulates thermoresponsive growth through TOC1 and PRR5. Genetic analyses reveal that ZTL regulates PIF4 activity as well as PIF4 expression. In Arabidopsis thaliana, ztl mutants exhibit highly accumulated TOC1 and PRR5 and unresponsive expression of PIF4 target genes under exposure to high temperatures. Mutations in TOC1 and PRR5 restore thermoactivation of PIF4 target genes and thermoresponsive growth in ztl mutants. We also show that the molecular chaperone heat‐shock protein 90 promotes thermoresponsive growth through the ZTL‐TOC1/PRR5 signaling module. Further, we show that ZTL protein stability is increased at high temperatures. Taken together, our results demonstrate that ZTL‐mediated degradation of TOC1 and PRR5 enhances the sensitivity of hypocotyl growth to high temperatures. Summary statement: A central clock component, ZEITLUPE (ZTL), is known to regulate thermoresponsive growth in Arabidopsis thaliana. However, its underlying mechanism remains poorly understood. This study demonstrates that ZTL regulates PIF4 activity as well as PIF4 expression through TOC1 (TIMING OF CAB EXPRESSION1) and PRR5 (PSEUDO‐RESPONSE REGULATORS5), thereby promoting thermomorphogenesis. [ABSTRACT FROM AUTHOR]

Auxin is well known to stimulate coleoptile elongation and rapid seedling growth in the air. However, its role in regulating rice germination and seedling establishment under submergence is largely unknown. Previous studies revealed that excessive levels of indole‐3‐acetic acid(IAA) frequently cause the inhibition of plant growth and development. In this study, the high‐level accumulation of endogenous IAA is observed under dark submergence, stimulating rice coleoptile elongation but limiting the root and primary leaf growth during anaerobic germination (AG). We found that oxygen and light can reduce IAA levels, promote the seedling establishment and enhance rice AG tolerance. miRNA microarray profiling and RNA gel blot analysis results show that the expression of miR167 is negatively regulated by submergence; it subsequently modulates the accumulation of free IAA through the miR167‐ARF‐GH3 pathway. The OsGH3‐8 encodes an IAA‐amido synthetase that functions to prevent free IAA accumulation. Reduced miR167 levels or overexpressing OsGH3‐8 increase auxin metabolism, reduce endogenous levels of free IAA and enhance rice AG tolerance. Our studies reveal that poor seed germination and seedling growth inhibition resulting from excessive IAA accumulation would cause intolerance to submergence in rice, suggesting that a certain threshold level of auxin is essential for rice AG tolerance. Summary statement: Excessive auxin accumulation negatively affects rice AG. Conversely, oxygen and light significantly reduce IAA levels, promote the seedling establishment during AG, and submergence‐repressed miR167 reduces IAA levels through miR167‐ARF‐GH3 pathway and enhances AG tolerance. [ABSTRACT FROM AUTHOR]

Leaf growth relies on photosynthesis and hydraulics to provide carbohydrates and expansion power; in turn, leaves intercept light and construct organism systems for functioning. Under potassium (K) deficiency stress, leaf area, photosynthesis and hydraulics are all affected by alterations in leaf structure. However, the connection between changes in leaf growth and function caused by the structure under K regulation is unclear. Consequently, the leaf hydraulic conductance (Kleaf) and photosynthetic rate (A) combined with leaf anatomical characteristics of Brassica napus were continuously observed during leaf growth under different K supply levels. The results showed that Kleaf and A decreased simultaneously after leaf area with the increasing K deficiency stress. K deficiency significantly increased longitudinal mesophyll cell investment, leading to a reduced volume fraction of intercellular air‐space (fias) and decreased leaf expansion rate. Furthermore, reduced fias decreased mesophyll and chloroplast surfaces exposed to intercellular airspace and gas phase H2O transport, which induced coordinated changes in CO2 mesophyll conductance and hydraulic conductance in extra‐xylem pathways. Adequate K supply facilitated higher fias through smaller palisade tissue cell density (loose mesophyll cell arrangement) and smaller spongy tissue cell size, which coordinated CO2 and H2O conductance and promoted leaf area expansion. [ABSTRACT FROM AUTHOR]

Plants have evolved to grow under prominently fluctuating environmental conditions. In experiments under controlled conditions, temperature is often set to artificial, binary regimes with constant values at day and at night. This study investigated how such a diel (24 hr) temperature regime affects leaf growth, carbohydrate metabolism and gene expression, compared to a temperature regime with a field‐like gradual increase and decline throughout 24 hr. Soybean (Glycine max) was grown under two contrasting diel temperature treatments. Leaf growth was measured in high temporal resolution. Periodical measurements were performed of carbohydrate concentrations, carbon isotopes as well as the transcriptome by RNA sequencing. Leaf growth activity peaked at different times under the two treatments, which cannot be explained intuitively. Under field‐like temperature conditions, leaf growth followed temperature and peaked in the afternoon, whereas in the binary temperature regime, growth increased at night and decreased during daytime. Differential gene expression data suggest that a synchronization of cell division activity seems to be evoked in the binary temperature regime. Overall, the results show that the coordination of a wide range of metabolic processes is markedly affected by the diel variation of temperature, which emphasizes the importance of realistic environmental settings in controlled condition experiments. Dynamically changing temperature affects soybean leaf growth and regulatory mechanisms in non‐intuitive ways. The results may help improving knowledge transfer between experiments performed under controlled conditions and experiments performed in the field. [ABSTRACT FROM AUTHOR]

To understand the growth response to drought, we performed a proteomics study in the leaf growth zone of maize (Zea mays L.) seedlings and functionally characterized the role of starch biosynthesis in the regulation of growth, photosynthesis and antioxidant capacity, using the shrunken-2 mutant (sh2), defective in ADP-glucose pyrophosphorylase. Drought altered the abundance of 284 proteins overrepresented for photosynthesis, amino acid, sugar and starch metabolism, and redox-regulation. Changes in protein levels correlated with enzyme activities (increased ATP synthase, cysteine synthase, starch synthase, RuBisCo, peroxiredoxin, glutaredoxin, thioredoxin and decreased triosephosphate isomerase, ferredoxin, cellulose synthase activities, respectively) and metabolite concentrations (increased ATP, cysteine, glycine, serine, starch, proline and decreased cellulose levels). The sh2 mutant showed a reduced increase of starch levels under drought conditions, leading to soluble sugar starvation at the end of the night and correlating with an inhibition of leaf growth rates. Increased RuBisCo activity and pigment concentrations observed in WT, in response to drought, were lacking in the mutant, which suffered more oxidative damage and recovered more slowly after re-watering. These results demonstrate that starch biosynthesis contributes to maintaining leaf growth under drought stress and facilitates enhanced carbon acquisition upon recovery. [ABSTRACT FROM AUTHOR]

Light is a key limiting factor of plant growth and development under the canopy. Specific light signals, such as a low ratio of red : far‐red (R:FR) light, trigger the shade avoidance response, which affects hypocotyl, stem, and leaf growth. Although multiple components mediating shade avoidance responses have been identified in the past few decades, the underlying regulatory mechanism remains unclear. In this study, we found that the far‐red elongated hypocotyls 3 (fhy3) mutant exhibited longer hypocotyls and increased expression levels of core shade avoidance response genes under low R:FR shade conditions compared with the wild type No‐0, suggesting that FHY3 negatively regulates shade avoidance responses. Yeast one‐hybrid, chromatin immunoprecipitation, and RT‐qPCR assays revealed that FHY3 directly binds to the promoters and gene body of PHYTOCHROME RAPIDLY REGULATED1 (PAR1) and PAR2 and activates their expression to inhibit shade responses. Furthermore, the overexpression of PAR1 or PAR2 rescued the enhanced shade avoidance responses of fhy3, indicating that both genes are direct downstream targets of FHY3 that mediate shade avoidance responses. Our findings demonstrate that the light‐signalling protein FHY3 positively regulates the transcription of PAR1 and PAR2, which encode two key negative regulators of shade avoidance responses, thus repressing plant responses to shade signals. Our findings demonstrate that the light‐signalling protein FHY3 positively regulates the transcription of PAR1 and PAR2, which encode two key negative regulators of shade avoidance responses, thus repressing plant responses to shade signals. [ABSTRACT FROM AUTHOR]

Basal branching in grasses, or tillering, is an important trait determining both form and function of crops. Although similarities exist between eudicot and grass branching programs, one notable difference is that the tiller buds of grasses are covered by the subtending leaf, whereas eudicot buds are typically unconstrained. The current study shows that contact with the leaf sheath represses sorghum bud growth by providing a mechanical signal that cues the bud to refrain from rapid growth. Leaf removal resulted in massive reprogramming of the bud transcriptome that included signatures of epigenetic modifications and also implicated several hormones in the response. Bud abscisic acid transiently increased, then decreased following leaf removal relative to controls, and abscisic acid was necessary to repress bud growth in the presence of the leaf. Jasmonic acid (JA) levels and signalling increased in buds following leaf removal. Remarkably, application of JA to buds in situ promoted growth. The repression of bud growth by leaf contact shares characteristics of thigmomorphogenic responses in other systems, including the involvement of JA, though the JA effect is opposite. The repression of bud growth by leaf contact may represent a mechanism to time tillering to an appropriate developmental stage of the plant. Sorghum tiller buds are repressed by mechanical signals generated by contact with the overlying leaf sheaths that alter bud‐local signalling by the phytohormones abscisic acid and jasmonic acid. [ABSTRACT FROM AUTHOR]

High‐resolution leaf growth is rarely studied despite its importance as a metric for plant performance and resource use efficiency. This is in part due to methodological challenges. Here, we present a method for in situ leaf growth measurements in a natural environment. We measured instantaneous leaf growth on a mature Avicennia marina subsp. australasica tree over several weeks. We measured leaf expansion by taking time‐lapse images and analysing them using marker tracking software. A custom‐made instrument was designed to enable long‐term field studies. We detected a distinct diel growth pattern with leaf area shrinkage in the morning and leaf expansion in the afternoon and at night. On average, the observed daily shrinkage was 37% of the net growth. Most of the net growth occurred at night. Diel leaf area shrinkage and recovery continued after growth cessation. The amount of daily growth was negatively correlated with shrinkage, and instantaneous leaf growth and shrinkage were correlated with changes in leaf turgor. We conclude that, at least in this tree species, instantaneous leaf growth patterns are very strongly linked to, and most likely driven by, leaf water relations, suggesting decoupling of short‐term growth patterns from carbon assimilation. High‐resolution leaf growth measurements in Avicennia marina showed leaf shrinkage in the morning and expansion mainly during the night, and a strong correlation of leaf area change with turgor, suggesting that growth was driven by plant hydraulic status rather than by the rate of carbon uptake. [ABSTRACT FROM AUTHOR]

Abstract: Research in maize is often performed using inbred lines that can be readily transformed, such as B104. However, because the B104 line flowers late, the kernels do not always mature before the end of the growing season, hampering routine seed yield evaluations of biotech traits introduced in B104 at many geographical locations. Therefore, we generated five hybrids by crossing B104 with the early‐flowering inbred lines CML91, F7, H99, Mo17, and W153R and showed in three consecutive years that the hybrid lines proved to be suitable to evaluate seed yield under field conditions in a temperate climate. By assessing the two main processes driving maize leaf growth, being rate of growth (leaf elongation rate or LER) and the duration of growth (leaf elongation duration or LED) in this panel of hybrids, we showed that leaf growth heterosis was mainly the result of increased LER and not or to a lesser extent of LED. Ectopic expression of the transgenes GA20‐oxidase (GA20‐OX) and PLASTOCHRON1 (PLA1), known to stimulate the LER and LED, respectively, in the hybrids showed that leaf length heterosis can be stimulated by increased LER, but not by LED, indicating that LER rather than LED is the target for enhancing leaf growth heterosis. [ABSTRACT FROM AUTHOR]

Strobilurins are an important class of agrochemical fungicides used throughout the world on a wide variety of crops as protection against fungal pathogens. In addition to this protective role, they are reported to also positively influence plant physiology. In this study, we analysed the effect of Stroby® WG, a commercially available fungicide consisting of 50% (w/w) kresoxim-methyl (KM) as active strobilurin compound, on Arabidopsis leaf growth. Treatment of seedlings with Stroby resulted in larger leaves due to an increase in cell number. Transcriptome analysis of Stroby-treated rosettes demonstrated an increased expression of genes involved in redox homeostasis, iron metabolism and sugar transport. Stroby treatment strongly induced the expression of the subgroup Ib basic helix-loop-helix (bHLH) transcription factors, which have a role in iron homeostasis under iron-limiting conditions. Single loss-of-function mutants of three bHLHs and their triple bhlh039, bhlh100 and bhlh101 mutant did not respond to Stroby treatment. Although iron and sucrose content was not affected, nitric oxide (NO) levels and nitrate reductase (NR) activity were significantly increased in Stroby-treated rosettes as compared with control plants. In conclusion, we suggest that the Stroby-mediated effects on growth depend on the increased expression of the subgroup Ib bHLHs and higher NO levels. [ABSTRACT FROM AUTHOR]

Leaf growth is controlled by various internal and external factors. Leaves of dicotyledonous plants show pronounced diel (24 h) growth patterns that are controlled by the circadian clock. To date, it is still uncertain whether diel leaf growth patterns remain constant throughout the development of a plant. In this study, we followed growth from the primary leaves to leaflets of the seventh trifoliate leaf of soybean ( G lycine max) on the same plants with a recently developed imaging-based method under controlled conditions and at a high temporal resolution. We found that all leaflets displayed a consistent diel growth pattern with maximum growth towards the end of the night. In some leaves, growth maxima occurred somewhat later - at dawn - as long as the leaves were still in a very early developmental stage. Yet, overall, diel growth patterns of leaves from different positions within the canopy were highly synchronous. Therefore, the diel growth pattern of any leaf at a given point in time is representative for the overall diel growth pattern of the plant leaf canopy and a deviation from the normal diel growth pattern can indicate that the plant is currently facing stress. [ABSTRACT FROM AUTHOR]

We have tested to what extent the growth ability of several organs of maize share a common genetic control. Every night, leaf elongation rate reaches a maximum value ( LERmax) that has a high heritability, is repeatable between experiments and is correlated with final leaf length. Firstly, we summarized quantitative trait loci (QTLs) of LERmax and of leaf length in three mapping populations. Among the 14 consensus QTLs ( cQTLs) of leaf length, seven co-located with cQTLs of LERmax with consistent allelic effects. Nine cQTLs of LERmax (4% of the genome) were highly reliable and confirmed by introgression lines. We then compared these QTLs with those affecting the growths of leaves, shoots, roots or young reproductive organs, detected with the same mapping populations in three field experiments or in literature datasets. Five of the nine most reliable cQTLs of LERmax co-located with QTLs involved in the growth of other organs (but not in flowering time) with consistent allelic effects. Reciprocally, two-thirds of the 20 QTLs of growth of different organs co-located with cQTLs of LERmax. Hence, LERmax, as determined in a phenotyping platform, is an indicator of the growth ability of other organs of the plant in controlled or in-field conditions. [ABSTRACT FROM AUTHOR]

ABSTRACT Methionine (Met) in proteins can be oxidized to two diastereoisomers of methionine sulfoxide, Met- S-O and Met- R-O, which are reduced back to Met by two types of methionine sulfoxide reductases (MSRs), A and B, respectively. MSRs are generally supplied with reducing power by thioredoxins. Plants are characterized by a large number of thioredoxin isoforms, but those providing electrons to MSRs in vivo are not known. Three MSR isoforms, MSRA4, MSRB1 and MSRB2, are present in Arabidopsis thaliana chloroplasts. Under conditions of high light and long photoperiod, plants knockdown for each plastidial MSR type or for both display reduced growth. In contrast, overexpression of plastidial MSRBs is not associated with beneficial effects in terms of growth under high light. To identify the physiological reductants for plastidial MSRs, we analyzed a series of mutants deficient for thioredoxins f, m, x or y. We show that mutant lines lacking both thioredoxins y1 and y2 or only thioredoxin y2 specifically display a significantly reduced leaf MSR capacity (-25%) and growth characteristics under high light, related to those of plants lacking plastidial MSRs. We propose that thioredoxin y2 plays a physiological function in protein repair mechanisms as an electron donor to plastidial MSRs in photosynthetic organs. [ABSTRACT FROM AUTHOR]

ABSTRACT Functions of α- and β-branch carotenoids in whole-plant acclimation to photo-oxidative stress were studied in Arabidopsis thaliana wild-type (wt) and carotenoid mutants, lutein deficient ( lut2, lut5), non- photochemical quenching1 ( npq1) and suppressor of zeaxanthin- less1 ( szl1) npq1 double mutant. Photo-oxidative stress was applied by exposing plants to sunflecks. The sunflecks caused reduction of chlorophyll content in all plants, but more severely in those having high α- to β-branch carotenoid composition ( α/ β-ratio) ( lut5, szl1npq1). While this did not alter carotenoid composition in wt or lut2, which accumulates only β-branch carotenoids, increased xanthophyll levels were found in the mutants with high α/ β-ratios ( lut5, szl1npq1) or without xanthophyll-cycle operation ( npq1, szl1npq1). The PsbS protein content increased in all sunfleck plants but lut2. These changes were accompanied by no change ( npq1, szl1npq1) or enhanced capacity (wt, lut5) of NPQ. Leaf mass per area increased in lut2, but decreased in wt and lut5 that showed increased NPQ. The sunflecks decelerated primary root growth in wt and npq1 having normal α/ β-ratios, but suppressed lateral root formation in lut5 and szl1npq1 having high α/ β-ratios. The results highlight the importance of proper regulation of the α- and β-branch carotenoid pathways for whole-plant acclimation, not only leaf photoprotection, under photo-oxidative stress. [ABSTRACT FROM AUTHOR]

ABSTRACT 13C discrimination between atmosphere and bulk leaf matter (Δ13 Clb) is frequently used as a proxy for transpiration efficiency ( TE). Nevertheless, its relevance is challenged due to: (1) potential deviations from the theoretical discrimination model, and (2) complex time integration and upscaling from leaf to whole plant. Six hybrid genotypes of Populus deltoides× nigra genotypes were grown in climate chambers and tested for whole-plant TE (i.e. accumulated biomass/water transpired). Net CO2 assimilation rates ( A) and stomatal conductance ( gs) were recorded in parallel to: (1) 13C in leaf bulk material ( δ13 Clb) and in soluble sugars ( δ13 Css) and (2) 18O in leaf water and bulk leaf material. Genotypic means of δ13 Clb and δ13 Css were tightly correlated. Discrimination between atmosphere and soluble sugars was correlated with daily intrinsic TE at leaf level (daily mean A/ gs), and with whole-plant TE. Finally, gs was positively correlated to 18O enrichment of bulk matter or water of leaves at individual level, but not at genotype level. We conclude that Δ13 Clb captures efficiently the genetic variability of whole-plant TE in poplar. Nevertheless, scaling from leaf level to whole-plant TE requires to take into account water losses and respiration independent of photosynthesis, which remain poorly documented. [ABSTRACT FROM AUTHOR]

ABSTRACT Light and soil water content affect leaf surface area expansion through modifications in epidermal cell numbers and area, while effects on leaf thickness and mesophyll cell volumes are far less documented. Here, three-dimensional imaging was applied in a study of Arabidopsis thaliana leaf growth to determine leaf thickness and the cellular organization of mesophyll tissues under moderate soil water deficit and two cumulative light conditions. In contrast to surface area, thickness was highly conserved in response to water deficit under both low and high cumulative light regimes. Unlike epidermal and palisade mesophyll tissues, no reductions in cell number were observed in the spongy mesophyll; cells had rather changed in volume and shape. Furthermore, leaf features of a selection of genotypes affected in leaf functioning were analysed. The low-starch mutant pgm had very thick leaves because of unusually large palisade mesophyll cells, together with high levels of photosynthesis and stomatal conductance. By means of an open stomata mutant and a 9- cis-epoxycarotenoid dioxygenase overexpressor, it was shown that stomatal conductance does not necessarily have a major impact on leaf dimensions and cellular organization, pointing to additional mechanisms for the control of CO2 diffusion under high and low stomatal conductance, respectively. [ABSTRACT FROM AUTHOR]

ABSTRACT The stable hydrogen isotope ratios ( δD) of leaf wax n-alkanes record valuable information on plant and ecosystem water relations. It remains, however, unknown if leaf wax n-alkane δD values record only environmental variation during the brief period of time of leaf growth or if leaf wax n-alkane δD values are affected by environmental variability throughout the entire lifespan of a leaf. To resolve these uncertainties, we irrigated Populus trichocarpa trees with a pulse of deuterium-enriched water and used compound-specific stable hydrogen isotope analyses to test if the applied tracer could be recovered from leaf wax n-alkanes of leaves that were at different stages of their development during the tracer application. Our experiment revealed that only leaf wax n-alkanes from leaves that had developed during the time of the tracer application were affected, while leaves that were already fully matured at the time of the tracer application were not. We conclude from our study that under controlled environmental conditions, leaf wax n-alkanes are synthesized only early in the ontogeny of a leaf. Our experiment has implications for the interpretation of leaf wax n-alkane δD values in an environmental context, as it suggests that these compounds record only a brief period of the environmental variability that a leaf experiences throughout its life. [ABSTRACT FROM AUTHOR]

Rice is known to be sensitive to soil water deficit and evaporative demand, with a greatest sensitivity of lowland-adapted genotypes. We have analysed the responses of plant water relations and of leaf elongation rate (LER) to soil water status and evaporative demand in seven rice genotypes belonging to different species, subspecies, either upland- or lowland-adapted. In the considered range of soil water potential (0 to −0.6 MPa), stomatal conductance was controlled in such a way that the daytime leaf water potential was similar in well-watered, droughted or flooded conditions (isohydric behaviour). A low sensitivity of LER to evaporative demand was observed in the same three conditions, with small differences between genotypes and lower sensitivity than in maize. The sensitivity of LER to soil water deficit was similar to that of maize. A tendency towards lower sensitivities was observed in upland than lowland genotypes but with smaller differences than expected. We conclude that leaf water status and leaf elongation of rice are not particularly sensitive to water deficit. The main origin of drought sensitivity in rice may be its poor root system, whose effect was alleviated in the study presented here by growing plants in pots whose soil was entirely colonized by roots of all genotypes. [ABSTRACT FROM AUTHOR]

A precise knowledge of the temporal and spatial distributions of cell division and tissue expansion is essential for appropriate leaf sampling in omics studies and for analyses of plant–environment relations. Elongating leaves of rice were studied during their whole development for elongation rate, distribution of cell length, cell production rate and spatial distribution of growth in the leaf. In seven genotypes, the pattern of leaf elongation rate followed three phases: (1) an exponential increase before leaf appearance; (2) a short phase (2–4 d at 20 °C) with a stable leaf elongation rate around leaf appearance; and (3) a phase of 8–10 d with a progressive decrease in elongation rate. The profile of cell length along the leaf changed with time during the first and last phases, but was time invariant around appearance. We propose a method adapted to non-steady elongation based on anatomical measurements, which was successfully tested by comparing it with the pricking method. It allowed analysis of the change with time in the spatial distribution of growth from initiation to end of leaf growth. The length of leaf zones with cell division and tissue elongation varied with time, with maximums of 21 and 60 mm respectively around leaf appearance. [ABSTRACT FROM AUTHOR]

The leaf is a multi-scale dynamic unit that is determined by mechanisms at different organizational scales (cell, tissue, whole leaf and whole plant) and affected by both internal (genotype) and external (environmental) determinisms. The recent development of phenotyping platforms and imaging techniques provides new insights into the temporal and spatial patterns of leaf growth as affected by those determinisms. Conclusions about the overriding mechanisms often depend on the considered organizational scale and of time resolution which varies from minutes to several weeks. Analyses of leaf growth responses to environmental conditions have revealed robust emerging properties at whole plant or whole leaf scales. They have highlighted that the control of individual leaf expansion is more complex than merely the sum of cellular processes, and the control at the whole plant level is more complex than the sum of individual leaf expansions. However, in many cases, the integrated leaf-growth variable can be simplified to a limited set of underlying variables to be measured for comparative analyses of leaf growth or modelling purposes. [ABSTRACT FROM AUTHOR]

Caterpillars and spider mites are herbivores with different feeding mechanisms. Spider mites feed on the cell content via stylets, while caterpillars, as chewing herbivores, remove larger amounts of photosynthetically active tissue. We investigated local and systemic effects of short-term caterpillar and spider mite herbivory on cotton in terms of primary metabolism and growth processes. After short-term caterpillar feeding, leaf growth and water content were decreased in damaged leaves. The glutamate/glutamine ratio increased and other free amino acids were also affected. In contrast, mild spider mite infestation did not affect leaf growth or amino acid composition, but led to an increase in total nitrogen and sucrose concentrations. Both herbivores induced locally increased dark respiration, suggesting an increased mobilization of storage compounds potentially available for synthesis of defensive substances, but did not affect assimilation and transpiration. Systemically induced leaves were not significantly affected by the treatments performed in this study. The results show that cotton plants do not compensate the loss of photosynthetic tissue with higher photosynthetic efficiency of the remaining tissue. However, early plant responses to different herbivores leave their signature in primary metabolism, affecting leaf growth. Changes in amino acid concentrations, total nitrogen and sucrose content may affect subsequent herbivore performance. [ABSTRACT FROM AUTHOR]

As ribulose 1·5-bisphosphate carboxylase/oxygenase (Rubisco) activity limits light-saturated photosynthesis under present atmospheric condition, the effects of an overexpression of RBCSon Rubisco content and photosynthesis were examined in the leaves at different positions in rice ( Oryza sativa L.). Rubisco content in the transformant was significantly greater in the uppermost, fully expanded leaves but decreased to levels similar to those in wild-type plants in the lower leaves. The mRNA levels of total RBCS and rbcL in these leaves were much less than those in the expanding leaves, where Rubisco synthesis is active, suggesting commensurately low level of synthesis. Although the activation state of Rubisco was lower in the uppermost, fully expanded leaves of the transformant, it recovered to its full level in the lower leaves. As a result, the photosynthetic rate did not differ in leaves at the same position between the transformant and the wild type. Similarly, whole plant biomass did not differ between these genotypes. Thus, we conclude that although the overexpression of RBCS led to an enhancement of Rubisco protein content in the uppermost, fully expanded leaves, it does not result in increased photosynthetic rates or plant biomass, because of an apparent down-regulation in its activation state. [ABSTRACT FROM AUTHOR]

Nitrogen deficiency severely inhibits leaf growth. This response was analysed at the cellular level by growing Lolium perenne L. under 7.5 mm (high) or 1 mm (low) nitrate supply, and performing a kinematic analysis to assess the effect of nitrogen status on cell proliferation and cell growth in the leaf blade epidermis. Low nitrogen supply reduced leaf elongation rate (LER) by 43% through a similar decrease in the cell production rate and final cell length. The former was entirely because of a decreased average cell division rate (0.023 versus 0.032 h−1) and thus longer cell cycle duration (30 versus 22 h). Nitrogen status did not affect the number of division cycles of the initial cell's progeny (5.7), and accordingly the meristematic cell number (53). Meristematic cell length was unaffected by nitrogen deficiency, implying that the division and mitotic growth rates were equally impaired. The shorter mature cell length arose from a considerably reduced post-mitotic growth rate (0.033 versus 0.049 h−1). But, nitrogen stress did not affect the position where elongation stopped, and increased cell elongation duration. In conclusion, nitrogen deficiency limited leaf growth by increasing the cell cycle duration and decreasing mitotic and post-mitotic elongation rates, delaying cell maturation. [ABSTRACT FROM AUTHOR]

Limited information is available on the genetic variation and control for plant growth response to elevated CO2 (e[CO2]). Such information is necessary to understand plant adaptation and evolution in future rising CO2. Here, quantitative trait loci (QTL) for leaf growth, development, quality and leaf senescence were determined in a tree pedigree – an F2 hybrid of Populus trichocarpa T. & G and Populus deltoides Marsh, following season-long exposure to either current day ambient carbon dioxide (a[CO2]) or e[CO2] at 600 µL L−1. Leaf growth and development differed between the grandparents such that P. trichocarpa showed greater response to e[CO2]. In the F2 generation, leaf development and quality traits including leaf area, leaf shape, epidermal cell area, and stomatal number, specific leaf area (SLA), and the phenology trait, canopy senescence index, were sensitive to e[CO2]. Sixty-nine QTL were mapped for the 19 traits of plants in a[CO2] while 60 QTL were mapped for plants in e[CO2]. The results suggest that although many QTL mapped to common positions in a[CO2] and e[CO2], confirming their importance in determining growth, there was also differential genetic control for a number of traits including leaf senescence. Candidate genes were shown to collocate to regions where response QTL mapped. This study is the first to identify candidate genes that may be important in determining plant adaptation to future high-CO2 world. [ABSTRACT FROM AUTHOR]

The role of abscisic acid (ABA) and its possible interaction with ethylene in mediating leaf elongation response to soil water deficit are a matter of controversy. To address this question, we used a set of maize genotypes with various levels of ABA either due to natural variability or to genetic transformation targeted on NCED/VP14, a key enzyme of ABA synthesis. The transgenic lines yielded less strong phenotypes than available mutants, making it possible to use them under normal growing conditions. We focused on leaf elongation during night periods in order to avoid the confounding effect of ABA on leaf water status. Our results suggest that over a wide range, internal ABA level (measured in both leaf extracts or xylem sap) has no clear effect on leaf elongation response to soil water deficit, except in the case of an antisense line presenting the strongest reduction in ABA accumulation that showed a slight maintenance of leaf elongation during water deficit. Leaf ethylene production rate was variable and not related to water deficit except in the ABA-deficient transgenic lines where it was increased by water deficit on average but not systematically. Moreover, variability in ethylene production rate was not linked to variability in elongation rate. Our results thus suggest that neither ABA nor ethylene seems to play a major role in the control of leaf elongation response to soil water deficit. [ABSTRACT FROM AUTHOR]

Limited information is available on the genetic variation and control for plant growth response to elevated CO2 (e[CO2]). Such information is necessary to understand plant adaptation and evolution in future rising CO2. Here, quantitative trait loci (QTL) for leaf growth, development, quality and leaf senescence were determined in a tree pedigree – an F2 hybrid of Populus trichocarpa T. & G and Populus deltoides Marsh, following season-long exposure to either current day ambient carbon dioxide (a[CO2]) or e[CO2] at 600 µL L−1. Leaf growth and development differed between the grandparents such that P. trichocarpa showed greater response to e[CO2]. In the F2 generation, leaf development and quality traits including leaf area, leaf shape, epidermal cell area, and stomatal number, specific leaf area (SLA), and the phenology trait, canopy senescence index, were sensitive to e[CO2]. Sixty-nine QTL were mapped for the 19 traits of plants in a[CO2] while 60 QTL were mapped for plants in e[CO2]. The results suggest that although many QTL mapped to common positions in a[CO2] and e[CO2], confirming their importance in determining growth, there was also differential genetic control for a number of traits including leaf senescence. Candidate genes were shown to collocate to regions where response QTL mapped. This study is the first to identify candidate genes that may be important in determining plant adaptation to future high-CO2 world. [ABSTRACT FROM AUTHOR]

To test whether the inhibition of leaf expansion by high evaporative demand is a result of hydraulic processes, we have followed both leaf elongation rate (LER) and cell turgor in leaves of maize plants either normally watered or in water-saturated soil in which hydraulic resistance at the soil–root interface was abolished. Cell turgor was measured in situ with a pressure probe in the elongating zone of the first and sixth leaves, and LERs of the same leaves were measured continuously with transducers or by following displacements of marks along the growing leaves. Both variables displayed spatial variations along the leaf and positively correlated within the elongating zone. Values peaked at mid-distance of this zone, where the response of turgor to evaporative demand was further dissected. High evaporative demand decreased both LER and turgor for at least 5 h, with dose-effect linear relations. This was observed in five genotypes with appreciable differences in turgor maintenance among genotypes. In contrast, the depressing effects of evaporative demand on both turgor and LER disappeared when the soil was saturated, thereby opposing a negligible resistance to water flow at the soil–root interface. These results suggest that the response of LER to evaporative demand has a hydraulic origin, enhanced by the resistance to water flux at the soil–root interface. They also suggest that turgor is not completely maintained under high evaporative demand, and may therefore contribute to the reductions in LER observed in non-saturated soils. [ABSTRACT FROM AUTHOR]

This study tested whether leaf elongation rate ( LER, mm h−1) and its components – average relative elemental growth rate ( REGRavg, mm mm−1 h−1) and leaf growth zone length ( LLGZ, mm) – are related to phosphorus (P) concentration in the growth zone ( PLGZ, mg P g−1 tissue water) of Lolium perenne L. cv. Condesa and whether such relationships are modified by the arbuscular mycorrhizal fungus (AMF) Glomus hoi. Mycorrhizal and non-mycorrhizal plants were grown at a range of P supply rates and analysed at either the same plant age or the same tiller size (defined by the length of the sheath of the youngest fully expanded leaf). Both improved P supply (up to 95%) and AMF (up to 21%) strongly increased LER. In tillers of even-aged plants, this was due to increased REGRavg and LLGZ. In even-sized tillers, it was exclusively due to increased REGRavg. REGRavg was strictly related to PLGZ ( r2 = 0.95) and independent of tiller size. Conversely, LLGZ strictly depended on tiller size ( r2 = 0.88) and not on PLGZ. Hence, P status affected leaf growth directly only through effects on relative tissue expansion rates. Symbiosis with AMF did not modify these relationships. Thus, no evidence for P status-independent effects of AMF on LER was found. [ABSTRACT FROM AUTHOR]

The effects of reduced SBPase activity on growth and development were examined in a set of transgenic tobacco plants produced using an antisense construct driven by the ribulose bisphosphate carboxylase, small subunit promoter. Photosynthetic carbon assimilation rates and carbohydrate levels in source leaves were decreased in the antisense plants. Growth rate and total shoot biomass were reduced in the SBPase antisense plants, even in plants where SBPase activity was reduced by only 25%. Floral biomass also decreased in response to reductions in SBPase activity and the onset of flowering was delayed by 5–10 d. This is the first demonstration of a link between reproductive biomass and reductions in Calvin cycle enzyme activity using antisense plants. Furthermore, unexpected changes in the growth and development of the antisense plants were evident. Small reductions in SBPase activity (above 50% wild type) resulted in shorter plants with only a small decrease in stem biomass and specific leaf area. In contrast, plants with larger reductions in SBPase activity had an increase in specific leaf area and attained heights similar to that of the wild-type plants but with a much reduced stem biomass, largely due to a decrease in xylem tissue. This bi-modal response of growth to reductions in SBPase activity has similarities to changes in leaf and stem anatomy and morphology that accompany light acclimation. [ABSTRACT FROM AUTHOR]

Leaf development is affected by both internal (genetic) and external (environmental) regulatory factors. The aim of this work was to investigate how leaf growth variables are related to one another in a range of environments. The leaf growth variables of wild-type Arabidopsis thaliana and leaf development mutants ( ang4, ron2-1, elo1, elo2 and elo4) were studied under different incident light treatments (light and shade). The leaves studied were altered in various leaf development variables, such as the duration of expansion, relative and absolute expansion rates, epidermal cell size, epidermal cell number and initiation rate. Final leaf area was correlated to maximal absolute leaf expansion rate and cell number, but not to duration of leaf expansion or cell size. These relationships were common to all studied genotypes and light conditions, suggesting that leaf size is determined early in development. In addition, the early variables involved in leaf development were correlated to one another, and initial relative expansion rate was negatively correlated to the duration of expansion. These relationships between the leaf development variables were used to construct a conceptual model of leaf size control. [ABSTRACT FROM AUTHOR]

The present authors have shown previously that both respiration rates and in vivo activities of the alternative oxidase (AOX) of leaves of Alocasia odora, a shade species, are lower than those in sun species, thereby optimizing energy production under limited light conditions (Noguchi et al., Australian Journal of Plant Physiology 28, 27–35, 2001). In the present study, mitochondria isolated from A. odora leaves were examined in order to investigate the biochemical basis for the differences in respiratory parameters. Alocasia odora and spinach plants were cultivated under both high and low light intensities, mitochondria were isolated from their leaves, and their respiratory properties compared. Mitochondrial content of leaf extracts from the two species was estimated using fumarase activities and antibody detection of porin (the voltage-dependent anion channel of the outer mitochondrial membrane). On a mitochondrial protein basis, spinach leaves showed higher capacities of the cytochrome pathway and cytochrome c oxidase (COX) than A. odora leaves. However, on a mitochondrial protein basis, A. odora showed higher capacities of AOX, which had a high affinity for ubiquinone when activated by pyruvate. Alocasia odora also had larger amounts of mitochondrial protein per leaf dry weight, even under severely shaded conditions, than spinach. Lower growth light intensity led to lower activities of most pathways and proteins tested in both species, especially glycine-dependent oxygen uptake. In the low light environment, most of the AOX protein in A. odora leaves was in its inactive, oxidized dimer form, but was converted to its reduced active form when plants were grown under high light. This shift may prevent over-reduction of the respiratory chain under photo-oxidative conditions. [ABSTRACT FROM AUTHOR]

Net fluxes of H+, K+ and Ca2+ ions from maize (Zea maysL.) isolated leaf segments were measured non-invasively using ion-selective vibrating microelectrodes (the MIFE technique). Leaf segments were isolated from the blade base, containing actively elongating cells (basal segments), and from non-growing tip regions (tip segments). Ion fluxes were measured in response to bright white light (2600 µmol m−2 s−1) from either the leaf segments or the underlying mesophyll (after stripping the epidermis). Fluxes measured from the mesophyll showed no significant difference between basal and tip regions. In leaf segments (epidermis attached), light-induced flux kinetics of all ions measured (H+, Ca2+ and K+) were strikingly different between the two regions. It appears that epidermal K+ fluxes are required to drive leaf expansion growth, whereas in the mesophyll light-induced K+ flux changes are likely to play a charge balancing role. Light-stimulated Ca2+ influx was not directly attributable either to leaf photosynthetic performance or to leaf expansion growth. It is concluded that light-induced ion flux changes are associated with both leaf growth and photosynthesis. [ABSTRACT FROM AUTHOR]

The tropical rainforest mesocosm within the Biosphere 2 Laboratory, a model system of some 110 species developed over 12 years under controlled environmental conditions, has been subjected to a series of comparable drought experiments during 2000–2002. In each study, the mesocosm was subjected to a 4–6 week drought, with well-defined rainfall events before and after the treatment. Ecosystem CO2 uptake rate (Aeco) declined 32% in response to the drought, with changes occurring within days and being reversible within weeks, even though the deeper soil layers did not become significantly drier and leaf-level water status of most large trees was not greatly affected. The reducedAeco during the drought reflected both morphological and physiological responses. It is estimated that the drought-induced 32% reduction ofAeco has three principal components: (1) leaf fall increased two-fold whereas leaf expansion growth of some canopy dominants declined to 60%, leading to a 10% decrease in foliage coverage of the canopy. This might be the main reason for the persistent reduction ofAeco after rewatering. (2) The maximum photosynthetic electron transport rate at high light intensities in remaining leaves was reduced to 71% for three of the four species measured, even though no chronic photo-inhibition occurred. (3) Stomata closed, leading to a reduced ecosystem water conductance to water vapour (33% of pre-drought values), which not only reduced ecosystem carbon uptake rate, but may also have implications for water and energy budgets of tropical ecosystems. Additionally, individual rainforest trees responded differently, expressing different levels of stress and stress avoiding mechanisms. This functional diversity renders the individual response heterogeneous and has fundamental implications to scale leaf level responses to ecosystem dynamics. [ABSTRACT FROM AUTHOR]

The stimulation of dicotyledonous leaf growth by light depends on increased H+ efflux, to acidify and loosen the cell walls, and is enhanced by K+ uptake. The role of K+ is generally considered to be osmotic for turgor maintenance. In coleoptiles, auxin-induced cell elongation and wall acidification depend on K+ uptake through tetraethylammonium (TEA)-sensitive channels (Claussen et al., Planta 201, 227–234, 1997), and auxin stimulates the expression of inward-rectifying K+ channels ( . The role of K+ in growing, leaf mesophyll cells has been investigated in the present study by measuring the consequences of blocking K+ uptake on several growth-related processes, including solute accumulation, apoplast acidification, and membrane polarization. The results show that light-stimulated growth and wall acidification of young tobacco leaves is dependent on K+ uptake. Light-stimulated growth is enhanced three-fold over dark levels with increasing external K+, and this effect is blocked by the K+ channel blockers, TEA, Ba++ and Cs+. Incubation in 10 m m TEA reduced light-stimulated growth and K+ uptake by 85%, and completely inhibited light-stimulated wall acidification and membrane polarization. Although K+ uptake is significantly reduced in the presence of TEA, solute accumulation is increased. We suggest that the primary role of K+ in light-stimulated leaf growth is to provide electrical counterbalance to H+ efflux, rather than to contribute to solute accumulation and turgor maintenance. [ABSTRACT FROM AUTHOR]

Grassland plants suffer regular defoliation, causing loss of photosynthetic activity and internal resources. Consequently, re-foliation may be substrate-limited. The present study was undertaken to test the hypothesis that decreased C import in leaf growth zones is (partially) compensated by: (i) mobilization of substrate within growth zones; and (ii) increased efficiency of substrate use in leaf area expansion; but (iii) that these processes depend on the C status of growth zones at defoliation. Mixtures of a C3 ( Lolium perenne L.) and a C4 grass ( Paspalum dilatatum Poir.) were grown at 15 °C (C3 dominance) and 23 °C (C4 dominance). Individual plants thus grew in contrasting (light and temperature) environments before being defoliated. Defoliation caused a drastic and immediate decrease in C import, but effects on leaf area expansion were buffered by biomass mobilization in the growth zone and increases in specific leaf area of produced tissue. Thus, over the first 2 d post-defoliation, the amount of leaf area produced per unit imported C increased by 39 to 102% depending on treatment. The magnitude of these buffering responses was correlated with the concentration of water soluble carbohydrates in the growth zone at defoliation. Similar responses were observed for N, although defoliation effects were smaller and delayed relative to those on C import. This study demonstrates refoliation is sustained by short-term mobilization of reserves within the growth zone and reduced costs of produced leaf area, but that these mechanisms depend on growth zone C status at defoliation. [ABSTRACT FROM AUTHOR]

The gravitational component of water potential contributes a standing 0.01 MPa m−1 to the xylem tension gradient in plants. In tall trees, this contribution can significantly reduce the water potential near the tree tops. The turgor of cells in buds and leaves is expected to decrease in direct proportion with leaf water potential along a height gradient unless osmotic adjustment occurs. The pressure–volume technique was used to characterize height-dependent variation in leaf tissue water relations and shoot growth characteristics in young and old Douglas-fir trees to determine the extent to which growth limitation with increasing height may be linked to the influence of the gravitational water potential gradient on leaf turgor. Values of leaf water potential ( Ψl), bulk osmotic potential at full and zero turgor, and other key tissue water relations characteristics were estimated on foliage obtained at 13.5 m near the tops of young (approximately 25-year-old) trees and at 34.7, 44.2 and 55.6 m in the crowns of old-growth (approximately 450-year-old) trees during portions of three consecutive growing seasons. The sampling periods coincided with bud swelling, expansion and maturation of new foliage. Vertical gradients of Ψl and pressure–volume analyses indicated that turgor decreased with increasing height, particularly during the late spring when vegetative buds began to swell. Vertical trends in branch elongation, leaf dimensions and leaf mass per area were consistent with increasing turgor limitation on shoot growth with increasing height. During the late spring (May), no osmotic adjustment to compensate for the gravitational gradient of Ψl was observed. By July, osmotic adjustment had occurred, but it was not sufficient to fully compensate for the vertical gradient of Ψl. In tall trees, the gravitational component of Ψl is superimposed on phenologically driven changes in leaf water relations characteristics, imposing potential constraints on turgor that may be indistinguishable from those associated with soil water deficits. [ABSTRACT FROM AUTHOR]

Abstract The primary leaf of wheat (Triticum aestivum L. cv Maris Huntsman) was used as a model system to examine how elevated ultraviolet-B (UV-B; λ = 280–320 nm) radiation affected growth. A reduction in the rate and duration of growth of the primary leaf, in response to UV-B, was the result of changes in both the rate and extent of cell division and elongation. UV-B reduced the proportion of mitotically active cells (mitotic index) and increased the time taken for cell division (cell doubling time). Thus the supply of cells into the elongation zone was reduced, and this, coupled to a reduction in the rate of elongation, resulted in reduced leaf growth. This analysis of the spatial distribution of growth provided a means of calculating the age of cells within the leaves. Cells of UV-B-treated leaves were found to age more quickly than those of the controls. This analysis will enable future studies to take account of age-related changes when interpreting the response of plants to any number of environmental stresses that affect leaf development. [ABSTRACT FROM AUTHOR]

ABSTRACTLeaves expand to intercept light for photosynthesis, to take up carbon dioxide, and to transpire water for cooling and circulation. The extent to which they expand is determined partly by genetic constraints, and partly by environmental conditions signalling the plant to expand more or less leaf surface area. Leaves have evolved sophisticated sensory mechanisms for detecting these cues and responding with their own growth and function as well as influencing a variety of whole-plant behaviours. Leaf expansion itself is an integrating behaviour that ultimately determines canopy development and function, allocation of materials determining relative shoot: root volume, and the onset of reproduction. To understand leaf development, and in particular, how leaf expansion is regulated, we must know at the molecular level which biochemical processes accomplish cell growth. Physiological experimentation focusing on ion fluxes across the plasmamembrane is providing new molecular information on how light stimulates cell expansion in some dicotyledonous species. Genetic analyses in Arabidopsis, corn, and other species are rapidly generating a list of mutations and enzyme activities associated with leaf development and expansion. Combination of these approaches, using informed physiological interpretations of phenotypic variation will allow us in the future to identify genes encoding both the processes causing cell expansion, and the regulators of these events. [ABSTRACT FROM AUTHOR]

ABSTRACTThe relative significance of the use of stored or currently absorbed C for the growth of leaves or roots of Lolium perenne L. after defoliation was assessed by steady-state labelling of atmospheric CO2. Leaf growth for the first two days after defoliation was to a large extent dependent on the use of C reserves. The basal part of the elongating leaves was mainly new tissue and 91% of the C in this part of the leaf was derived from reserves assimilated prior to defoliation. However, half of the sucrose in the growth zone was produced from photosynthesis by the emerged leaves. Fructans that were initially present in elongating leaf bases were hydrolysed (loss of 93 to 100%) and the resulting fructose was found in the new leaf bases, suggesting that this pool may be used to support cell division and elongation. Despite a negative C balance at the whole-plant level, fructans were synthesized from sucrose that was translocated to the new leaf bases. After a regrowth period of 28 d, 45% of the C fixed before defoliation was still present in the root and leaf tissue and only 1% was incorporated in entirely new tissue. [ABSTRACT FROM AUTHOR]

The effects of nitrogen deprivation on leaf extension, cell numbers and epidermal cell size were followed in leaves of Ricinus communis L. The extent to which reductions in final cell number or final epidermal cell size contributed to the reduction in final leaf size depended on the developmental stage of the leaf at the time of N deprivation. In leaves which already had their full complement of cells (leaf 2), the reduction in final leaf size following nitrogen deprivation was associated with a reduction in final cell size. In leaves that were at earlier stages of development at the onset of N deprivation (leaves 3 and 4), the reduction in final leaf size was greater than in leaf 2. In these younger leaves, the final cell size was even smaller than in leaf 2, but the greatest contribution to reduced final leaf size was a reduction in the number of cells produced. This accounted for approximately 80% of the reduction in final leaf size in leaf 4. During leaf development, the contribution from different tissue layers to the total cell number changed. In the smallest leaf sizes, the contribution from upper and lower epidermis and spongy parenchyma was greater than that from palisade parenchyma. As the leaf size increased, cells in the palisade parenchyma continued to divide for longer than in the other layers. At final leaf size, the contribution from the different tissue layers to total cell number was the same for leaves 2, 3 and 4, irrespective of N treatment. In these final leaf structures, palisade parenchyma contributed 60% of the total cell number. Thus, although nitrogen deprivation affected leaf size variously through cell division and cell expansion, depending on leaf developmental stage at the time of nitrogen deprivation, the ratio of cell numbers and sizes in different tissue layers, at final leaf size, was unaffected. [ABSTRACT FROM AUTHOR]

We have tested whether the effects of temperature on sunflower leaf growth could be documented by using thermal time. The rates of leaf expansion and of cell division were analysed in leaves located at two positions on the stem, and a spatial analysis of expansion rate was carried out. Experiments were performed in growth chamber (stable conditions), in the field or in a greenhouse (fluctuating conditions). We compared three methods for characterizing the rate and the duration of expansion. Responses to leaf temperature were consistent only when expansion was characterized as a two-phase process — a period of exponential expansion (constant relative expansion rate, RER) followed by a decrease in RER. RER and relative cell division rate (RDR) responded linearly to temperature with a common response curve for all studied conditions. This response curve was also common to all studied zones within a leaf and to leaves at two positions on the stem. The reciprocals of the durations of the periods of exponential expansion, non-zero expansion and non-zero division were also linearly related to leaf temperature with common response curves in a given leaf zone. The x-intercepts of all these response curves and of the response curve of leaf initiation rate to temperature did not significantly differ in an analysis of covariance, with a common value of 4·8 °C. The expression of time in cumulative degree days, with a base temperature of 4·8 °C, resulted in a unique time course of RER and cell division rate regardless of temperature. These results suggest that a powerful ‘program’ of leaf development exists in a sunflower plant. [ABSTRACT FROM AUTHOR]

The osmotic and ion-specific components of salt-induced inhibition of leaf expansion growth were investigated in beans grown from 12 h to several days in either NaCl-containing solution cultures, an isosmotic concentrated macronutrient solution, or a vermiculite–compost mixture with low Na+ but high Cl– availability. Inhibition of leaf expansion and leaf ABA increase was more intense in the NaCl than in the isosmotic macronutrient treatment. Root Na+ was highly correlated to inhibition of leaf expansion and leaf or xylem sap ABA. When Na+ was sequestered in soil, salinized plants showed no reduction in leaf expansion or ABA increase, regardless of the presence of high leaf Cl– concentrations. Stomatal conductance exhibited an exponential relationship with the reciprocal value of xylem sap ABA. Our results indicate that an ion-specific effect caused by Na+ in roots may account for an ABA-mediated reponse of both stomatal closure and leaf expansion inhibition. [ABSTRACT FROM AUTHOR]

This study on expansive growth of the first leaf of maize has two goals: one is to determine how the sensitivity of growth to changes in water status varies with the initial water status of the leaf, and the other is to adapt the pressure-jump technique of Okamoto et al. (1989, Plant and Cell Physiology 30, 979–985), developed for studying growth of excised stem segments, for use on whole seedlings. Initial water status was varied by using: transpiring vs. non-transpiring conditions, seedlings differing in emerged leaf length and hence transpiring area, and root medium without mannitol vs. medium with added mannitol (to –0·3 MPa). The results show that growth changed with changes in plant water status when the water status was low, but was unaffected when water status was very high. A stepwise change in hydrostatic pressure on the root medium was quickly and fully transmitted to the base of the leaf. The increase in leaf elongation due to a pressure step of 0·025 MPa was negligible under conditions of high plant water status and became substantial under conditions of low water status. In adapting the pressure-jump method to the whole seedling, there was some loss of resolution, and the yield threshold Y of the Lockhart equation could not be estimated directly. Nonetheless, the data were suitable for the calculation of volumetric extensibility m and the estimation of growth effective turgor (turgor above Y). Extensibility was shown to increase 3- to 4-fold when leaf water status was reduced from the maximum to the point where elongation rate was halved, while growth effective turgor was calculated to diminish even more markedly. [ABSTRACT FROM AUTHOR]

The possibility that changes in the plasticity of expanding cell walls are involved in regulating early leaf growth responses to nutrientdeficiencies in monocot plants was investigated. Intact maize seedlings (Zea mays L.) which were hydroponically grown with their roots in low-nutrient solution (1 mol m-3 CaCl2) showed early inhibition of first-leaf growth, as compared with seedlings on complete nutrient solution. This early inhibition of leaf growth was not associated with reduced cell production. However, segmental elongation along the cell expansion zone at the base of the leaf and the lengths of mature epidermal cells were reduced by the low-nutrient treatment. Solute (osmotic) potentials in the expanding leaf tissues were unchanged. In contrast, low-nutrient treatments significantly altered leaf plasticity, i.e. the irreversible extension caused by applyinga small force in the direction of leaf growth. For example, in vivo plasticity decreased, along with leaf growth, after transfer of seedlings from complete nutrient solution to low-nutrient solution for 15 h. Conversely, in vivo plasticity increased, along with leaf growth, after transfer of plants previously grown on low-nutrient solution to complete nutrient solution for 15 h. The nutrient treatments also induced similar changes in the in vitro plasticity of the expanding leaf cell walls. There were no consistent changes in elasticity. Thus, reductions in the plasticity of expanding leaf cell walls appear to be involved in controlling the early inhibition of maize leaf growth by root imposition of nutrient stress. [ABSTRACT FROM AUTHOR]

Detached barley (Hordeum vulgare L) shoots, maintained at different air temperatures and VPDs, were fed ABA via the sub-crown internode in a leaf elongation assay. Analysis of variance of leaf elongation rate (LER) showed significant effects of temperature (T), fed [ABA] and the interaction T × [ABA]. However, the interaction became non-significant when LER was modelled against the [ABA] of the elongation zone, [EZ-ABA]. When detached barley shoots were fed sp from droughted maize (Zea mays L.) plants, sap [ABA] could not explain the growth inhibitory activity. Measurement of [EZ-ABA] accounted for this `unexplained' growth inhibition. The detached shoot experiments indicated that [EZ- ABA], and not xylem sap [ABA], was an appropriate explanatory variable to measure in droughted plants. However, ABA accumulation in the elongation zone could not explain a 35% growth reduction in intact droughted plants; thus we considered an interaction of water status and ABA. Using a coleoptile growth assay. we applied mild osmotic stresses (ψ=0 to -0.06 MPa) and 10-4 mol m-3 ABA. Individually, these treatments (lid not inhibit growth. However, osmotic stress and ABA applied together significantly reduced growth. This interaction may be an important mechanism in explaining leaf growth inhibition of droughted plants. [ABSTRACT FROM AUTHOR]

To evaluate the effect of different naturally occurring irradiation conditions on the sensitivity of bean (Phaseolus vulgaris cv. Label) to increased UV-B levels, plants were grown under six different light treatments. In the control series (at ambient levels of UV-B), UV-B and visible light were decreased in parallel, resulting in three different total irradiation treatments with the same UV-B/PAR ratio. A second series with a 15% increase in UV-B irradiation at each PAR level was used to investigate the effect of UV-B under the varying total irradiance levels. The different total irradiance levels resulted in large differences in total dry weight, specific leaf weight, photosynthesis-light response and pigment concentrations. Nevertheless, the 1S% increase in UV-B resulted In equal reductions in total dry weight (from 245 to 343%) and effective photosynthesis for all light levels. The accumulation of protective pigments in the primary bean leaves was strongly correlated to the total irradiance level (200% increase from the lowest to the highest light level), but was not influenced by increasing UV-B levels. As the UV-BIPAR ratio outside increases with decreasing total irradiance (when induced by cloud cover) this implies that low radiation levels are potentially dangerous to some plants, even though the UV-B levels may seem negligible. [ABSTRACT FROM AUTHOR]