Dynamics of leaf and root growth: endogenous control versus environmental impact

AIMS: Production of biomass and yield in natural and agronomic conditions depend on the endogenous growth capacity of plants and on the environmental conditions constraining it. Sink growth drives the competition for carbon, nutrients and water within the plant, and determines the structure of leaves and roots that supply resources to the plant later on. For their outstanding importance, analyses of internal growth mechanisms and of environmental impact on plant growth are long-standing topics in plant sciences. SCOPE: Recent technological developments have made it feasible to study the dynamics of plant growth in temporal and spatial scales that are relevant to link macroscopic growth with molecular control. These developments provided first insights into the truly dynamic interaction between environment and endogenous control of plant growth. CONCLUSIONS: Evidence is presented in this paper that the relative importance of endogenous control versus the impact of the dynamics of the environment depends on the frequency pattern of the environmental conditions to which the tissue is exposed. It can further be speculated that this is not only relevant within individual plants (hence leaves versus roots), but also crucial for the adaptation of plant species to the various dynamics of their environments. The following are discussed: mechanisms linking growth and concentrations of primary metabolites, and differences and homologies between spatial and temporal patterns of root and leaf growth with metabolite patterns.     

The photosynthetic response of tobacco plants overexpressing ice plant aquaporin McMIPB to a soil water deficit and high vapor pressure deficit

We investigated the photosynthetic capacity and plant growth of tobacco plants overexpressing ice plant (Mesembryanthemum crystallinum L.) aquaporin McMIPB under (1) a well-watered growth condition, (2) a well-watered and temporal higher vapor pressure deficit (VPD) condition, and (3) a soil water deficit growth condition to investigate the effect of McMIPB on photosynthetic responses under moderate soil and atmospheric humidity and water deficit conditions. Transgenic plants showed a significantly higher photosynthesis rate (by 48 %), higher mesophyll conductance (by 52 %), and enhanced growth under the well-watered growth condition than those of control plants. Decreases in the photosynthesis rate and stomatal conductance from ambient to higher VPD were slightly higher in transgenic plants than those in control plants. When plants were grown under the soil water deficit condition, decreases in the photosynthesis rate and stomatal conductance were less significant in transgenic plants than those in control plants. McMIPB is likely to work as a CO₂ transporter, as well as control the regulation of stomata to water deficits.     

Photosynthesis, N(2) fixation and taproot reserves during the cutting regrowth cycle of alfalfa under elevated CO(2) and temperature

Future climatic conditions, including rising atmospheric CO₂ and temperature may increase photosynthesis and, consequently, plant production. A larger knowledge of legume performance under the predicted growth conditions will be crucial for safeguarding crop management and extending the area under cultivation with these plants in the near future. N₂ fixation is a key process conditioning plant responsiveness to varying growth conditions. Moreover, it is likely to increase under future environments, due to the higher photosynthate availability, as a consequence of the higher growth rate under elevated CO₂. However, as described in the literature, photosynthesis performance is frequently down-regulated (acclimated) under long-term exposure to CO₂, especially when affected by stressful temperature and water availability conditions. As growth responses to elevated CO₂ are dependent on sink-source status, it is generally accepted that down-regulation occurs in situations with insufficient plant C sink capacity. Alfalfa management involves the cutting of shoots, which alters the source-sink relationship and thus the photosynthetic behaviour. As the growth rate decreases at the end of the pre-cut vegetative growth period, nodulated alfalfa plants show photosynthetic down-regulation, but during regrowth following defoliation, acclimation to elevated CO₂ disappears. The shoot harvest also leads to a drop in mineral N uptake and C translocation to the roots, resulting in a reduction in N₂ fixation due to the dependence on photosynthate supply to support nodule function. Therefore, the production of new shoots during the first days following cutting requires the utilization of reduced C and N compounds that have been stored previously in reserve organs. The stored reserves are mediated by phytohormones such as methyl jasmonate and abscisic acid and in situations where water stress reduces shoot production this potentially enables the enhancement of taproot protein levels in nodulated alfalfa, which may lead to these plants being in better condition in the following cut/regrowth cycle. Furthering our knowledge of legume performance under predicted climate change conditions will be crucial for the development of varieties with better adaptation that will achieve greater and more efficient production values. Furthermore, for this purpose it will be necessary to improve existing methodologies and create new ones for phenotype characterization. Such knowledge will provide key information for future plant breeding programs.     

Salicylic acid and photosynthesis: signalling and effects

Salicylic acid (SA) is a well-known signalling molecule playing a role in local and systemic acquired resistance against pathogens as well as in acclimation to certain abiotic stressors. As a stress-related signalling compound, it may directly or indirectly affect various physiological processes, including photosynthesis. The effects of exogenously applied SA on plant physiological processes under optimal environmental conditions are controversial. Several studies suggest that SA may have a positive effect on germination or plant growth in various plant species. However, SA may also act as a stress factor, having a negative influence on various physiological processes. Its mode of action depends greatly on several factors, such as the plant species, the environmental conditions (light, temperature, etc.) and the concentration. Exogenous SA may also alleviate the damaging effects of various stress factors, and this protection may also be manifested as higher photosynthetic capacity. Unfavourable environmental conditions have also been shown to increase the endogenous SA level in plants. Recent results strongly suggest that controlled SA levels are important in plants for optimal photosynthetic performance and for acclimation to changing environmental stimuli. The present review discusses the effects of exogenous and endogenous SA on the photosynthetic processes under optimal and stress conditions.     

植物对盐分空间不均匀分布的形态和生理响应研究进展

盐胁迫是干旱、半干旱地区以及灌溉土地主要的非生物胁迫,是影响农业生产的主要不利环境因子之一。随集约化灌溉农业的发展、水资源的缺乏、气候干旱带来的蒸发量的增加,土壤及地下水盐渍化程度不断增加。自然界中,土壤盐分在时空上呈不均匀分布。关于植物对均匀盐胁迫的响应研究报到较多,然而植物对不均匀盐胁迫的响应研究报道较少。分析了国内外植物适应不均匀盐胁迫的研究案例,从植物地上部分生长、地下部分生长、水分调节、光合作用以及离子调控等方面阐述植物适应盐分不均匀分布的生理机制,并提出展望。     

Triacontanol as a dynamic growth regulator for plants under diverse environmental conditions

Triacontanol (TRIA) being an endogenous plant growth regulator facilitates numerous plant metabolic activities leading to better growth and development. Moreover, TRIA plays essential roles in alleviating the stress-accrued alterations in crop plants via modulating the activation of the stress tolerance mechanisms. The present article critically focuses on the role of exogenously applied TRIA in morpho-physiology and biochemistry of plants for example, in terms of growth, photosynthesis, enzymatic activity, biofuel synthesis, yield and quality under normal and stressful conditions. This article also enlightens the mode of action of TRIA and its interaction with other phytohormones in regulating the physio-biochemical processes in counteracting the stress-induced damages in plants.

     

The effects of environmental heterogeneity on root growth and root/shoot partitioning

AIMS: The purpose of this Botanical Briefing is to stimulate reappraisal of root growth, root/shoot partitioning, and analysis of other aspects of plant growth under heterogeneous conditions. SCOPE: Until recently, most knowledge of plant growth was based upon experimental studies carried out under homogeneous conditions. Natural environments are heterogeneous at scales relevant to plants and in forms to which they can respond. Responses to environmental heterogeneity are often localized rather than plant-wide, and not always predictable from traditional optimization arguments or from knowledge of the ontogenetic trends of plants growing under homogeneous conditions. These responses can have substantial impacts, both locally and plant-wide, on patterns of resource allocation, and significant effects on whole-plant growth. Results from recent studies are presented to illustrate responses of plants, plant populations and plant communities to nutritionally heterogeneous conditions. CONCLUSIONS: Environmental heterogeneity is a constant presence in the natural world that significantly influences plant behaviour at a variety of levels of complexity. Failure to understand its effects on plants prevents us from fully exploiting aspects of plant behaviour that are only revealed under patchy conditions. More effort should be invested into analysis of the behaviour of plants under heterogeneous conditions.     

The Teaching of Photosynthesis in Secondary School: A History of the Science Approach

In this article we present a synthesis of the research affecting pupils' conceptions of photosynthesis and plant nutrition. The main false conceptions of the pupils identi?ed in this literature review are: that green plants ?nd their food in the soil; that water and mineral salts are suf?cient to the growth of a plant; the role of chlorophyll, where the transformation of luminous energy into chemical energy is never evoked; and air as a source of matter, which is never underlined. Secondly, we are going to see that several of these false conceptions have been developed during history. For example, the famous philosopher Aristotle (384–322 BC) thought that plants receive their food from the soil already elaborated. Several centuries later, the physician and chemist Van Helmont (1677–1644) added more precision to Aristotle’s conception while claiming that plants use only water for their growth. Finally, we will see that the analysis of the false theories developed during history will permit in a context of teaching to valorize the false conceptions of the pupils. Indeed, the history of sciences could incite a teacher to valorize his pupils’ false conceptions while considering them as an indication of difficulties that deserve particular pedagogical and didactic tools. The false conceptions constructed by the pupils don’t have to be ignored in a teaching context because they obey particular reasoning rules, sometimes similar to those that once guided some scientific steps. This view, drawing on false conceptions developed during history, gives a dynamic and human picture of the science very distant from the one sometimes carried on by dogmatic teaching.     

Is leaf-level photosynthesis related to plant success in a highly productive grassland?

We addressed the question: “Are short-term, leaf-level measurements of photosynthesis correlated with long-term patterns of plant success?” in a productive grassland where interspecific competitive interactions are important. To answer this question, seasonal patterns of leaf-level photosynthesis were measured in 27 tallgrass prairie species growing in sites that differed in species composition and productivity due to differences in fire history. Our specific goals were to assess the relationship between gas exchange under field conditions and success (defined as aerial plant cover) for a wide range of species, as well as for these species grouped as dominant and sub-dominant grasses, forbs, and woody plants. Because fire increases productivity and dominance by grasses in this system, we hypothesized that any relationship between photosynthesis and success would be strongest in annually burned sites. We also predicted that regardless of fire history, the dominant species (primarily C₄ grasses) would have higher photosynthetic rates than the less successful species (primarily C₃ grasses, forbs and woody plants). Because forbs and woody species are less abundant in annually burned sites, we expected that these species would have lower photosynthetic rates in annually burned than in infrequently burned sites. As expected, the dominant C₄?grasses had the highest cover on all sites, relative to?other growth forms, and they had the highest maximum and seasonally averaged photosynthetic rates (17.6 ± 0.42 μmol m^?2 s^?1). Woody species had the lowest average cover as well as the lowest average photosynthetic rates, with subdominant grasses and forbs intermediate in both cover and photosynthesis. Also as predicted, the highest overall photosynthetic rates were found on the most productive annually burned site. Perhaps most importantly, a positive relationship was found between leaf-level photosynthesis and cover for a core group of species when data were combined across all sites. These data support the hypothesis that higher instantaneous rates of leaf-level photosynthesis are indicative of long-term plant success in this grassland. However, in contrast to our predictions, the subdominant grasses, forbs and woody species on the annually burned site had higher photosynthetic rates than in the less frequently burned sites, even though their average cover was lower on annually burned sites, and hence they were less successful. The direct negative effect of fire on plant cover and species-specific differences in the availability of resources may explain why photosynthesis was high but cover was low in some growth forms in annually burned sites.     

Diurnal regulation of plant growth

Life occurs in an ever-changing environment. Some of the most striking and predictable changes are the daily rhythms of light and temperature. To cope with these rhythmic changes, plants use an endogenous circadian clock to adjust their growth and physiology to anticipate daily environmental changes. Most studies of circadian functions in plants have been performed under continuous conditions. However, in the natural environment, diurnal outputs result from complex interactions of endogenous circadian rhythms and external cues. Accumulated studies using the hypocotyl as a model for plant growth have shown that both light signalling and circadian clock mutants have growth defects, suggesting strong interactions between hypocotyl elongation, light signalling and the circadian clock. Here, we review evidence suggesting that light, plant hormones and the circadian clock all interact to control diurnal patterns of plant growth.     

Effects of N-supply on the rates of photosynthesis and shoot and root respiration of inherently fast- and slow-growing monocotyledonous species

Are there intrinsic differences in the rates of photosynthesis, shoot- and root-respiration between inherently fast- and slow-growing monocotyledons at high and low nitrogen supply? To analyze this question we grew 5 monocotyledons, widely differing in their inherent relative growth rate at high and low nitrogen supply in a growth room. Nitrate was exponentially added to the plants, enabling us to compare inherent differences in plant characteristics, without any effect of species differences in the ability to take up nutrients. At high nitrogen supply, the fast-growing species from productive habitats had a higher photosynthetic nitrogen use efficiency and rate of root respiration than the slow-growing ones from unproductive habitats. Only minor differences were observed in their rates of photosynthesis and shoot respiration per unit leaf area. At low nitrogen supply, the rates of photosynthesis and shoot- and root respiration decreased for all species, even though there were no longer any differences in these processes between inherently fast- and slow-growing species. The photosynthetic nitrogen use efficiency increased for all species, and no differences were found among species. Differences in the photosynthetic nitrogen use efficiency among species and nitrogen treatments are discussed in terms of the utilization of the photosynthetic apparatus, whereas differences in respiration rate are discussed in terms of the energy demand for growth, maintenance and ion uptake and their related specific respiratory energy costs. It is concluded that the relatively high abundance of slow-growing species compared to fast-growing ones in unproductive habitats is unlikely to be explained by differences in rates of photosynthesis and respiration or in photosynthetic nitrogen use efficiency.     

Impact of light limitation on seagrasses

Seagrass distribution is controlled by light availability, especially at the deepest edge of the meadow. Light attenuation due to both natural and anthropogenically-driven processes leads to reduced photosynthesis. Adaptation allows seagrasses to exist under these sub-optimal conditions. Understanding the minimum quantum requirements for growth (MQR) is revealed when light conditions are insufficient to maintain a positive carbon balance, leading to a decline in seagrass growth and distribution. Respiratory demands of photosynthetic and non-photosynthetic tissues strongly influence the carbon balance, as do resource allocations between above- and below-ground biomass. Seagrass light acclimation occurs on varying temporal scales, as well as across spatial scales, from the position along a single leaf blade to within the canopy and finally across the meadow. Leaf absorptance is regulated by factors such as pigment content, morphology and physical properties. Chlorophyll content and morphological characteristics of leaves such as leaf thickness change at the deepest edge. We present a series of conceptual models describing the factors driving the light climate and seagrass responses under current and future conditions, with special attention on the deepest edge of the meadow.     

Functional significance of triazine-herbicide resistance in defence of Senecio vulgaris against a rust fungus

We investigated the functional significance of plant performance (dry mass, photosynthesis) in plant defence (resistance and tolerance) against pathogen infection, and potential negative cross-resistance between herbicide resistance and plant defence against disease. We compared isonuclear triazine-herbicide-resistant (TR) and -susceptible (TS) biotypes of Senecio vulgaris, in the presence and absence of infection by the rust Puccinia lagenophorae. In a growth chamber study with two reduced irradiance levels, rust infection had a severe effect on plant performance with infected plants having 55% less dry mass and 54% reduced whole-plant photosynthesis than non-infected plants. The TR biotype was more susceptible (reduced resistance) to the pathogen, but the biotypes did not differ in their ability to compensate for rust infection (tolerance). TR plants were less productive than TS plants when grown non-shaded (ca. 10% full sunlight) but not when shaded (ca. 5% full sunlight). This is especially important for situations, where S. vulgaris grows under the crop canopy (e.g. in maize). Here, very low light levels might contribute to a numerical increase of TR relative to TS plants even when only occasionally treated with triazine. Whole-plant photosynthesis was reduced by 21% in TR plants as compared to the TS biotype, and by 59% in plants grown in the shaded as compared to the non-shaded treatment. When whole-plant photosynthesis values were corrected for the estimated leaf area of plants, we found no significant variation between biotypes, shade treatments or rust treatments. In experimental mixed TR:TS field populations, the proportion of TR plants decreased more rapidly in rust-infected populations than uninfected. This finding, together with the lower resistance in the TR than the TS biotype to the rust fungus observed in the growth chamber experiment, may indicate negative cross-resistance, which is a potential tool in the management of herbicide-resistant weeds.     

Regulation in Plant Stress Tolerance by a Potential Plant Growth Regulator, 5-Aminolevulinic Acid

Exogenous application of different plant growth regulators is a well-recognized strategy to alleviate stress-induced adverse effects on different crop plants by regulating a variety of physiobiochemical processes such as photosynthesis, chlorophyll biosynthesis, nutrient uptake, antioxidant metabolism, and protein synthesis, which are directly or indirectly involved in the mechanism of stress tolerance. Of various environmental factors, salinity, drought, and extreme temperature (low or high) considerably diminish plant growth and yield by modulating endogenous levels as well as signaling pathways of plant hormones. Of various plant hormones/regulators, a potential plant growth regulator, 5-aminolevulinic acid (ALA), is known to be effective in counteracting the injurious effects of various abiotic stresses in plants. Until now the mechanisms behind ALA regulation of growth under stress have not been fully elucidated. It is also not yet clear how far growth and yield in different crops can be promoted by exogenous application of ALA and whether this ALA-induced growth and yield promotion is cost-effective. Thus, in this review we discuss at length the effects of ALA in regulating growth and development in plants under a variety of abiotic stress conditions, including salinity, drought, and temperature stress. Furthermore, advances in the functional and regulatory interactions of this plant growth regulator with plant stress tolerance, as well as the effective mode of exogenous application of ALA in inducing stress tolerance in plants are also comprehensively discussed in this review. In the future, overaccumulation of ALA in plants through manipulation of gene(s) could enhance plant stress tolerance. Thus, genetic manipulation of plants with the goal of attaining increased synthesis/accumulation of ALA and hence improved stress tolerance under stress conditions is an important area for research.     

Phloem loading and its development related to plant evolution from trees to herbs

Summary Minor vein structure in various taxonomic groups was described in a previous paper (Gamalei 1989). Here, these results are used to correlate minor vein structure with plant evolutionary, ecological and growth form schemes. The following pattern emerges: reductive evolution from evergreen trees to annual herbs is accompanied by gradually increasing symplastic isolation of the mesophyll and the phloem. This evolutionary tendency is confirmed by the ecological spreading and life-form distribution of modern plants with different types of minor vein structure. The meaning of this phenomenon is discussed. Chilling sensitivity of plasmodesmal translocation is considered to be the main reason. It is suggested that phloem loading for assimilate transport is double-routed. The symplastic route is more ancient and more economical for loading. The apoplastic pathway becomes the main or the only route under unfavorable conditions. The existence of a symplast/apoplast regulatory loading mechanism is suggested. The two loading routes differ in their selectivity for products of photosynthesis which changes their symplast/apoplast ratio which, in turn, determines the composition of the sieve tube exudate. The latter will influence growth and morphogenesis. Correlated changes of structure and function related to photosynthesis, loading, translocation and growth, are analysed with respect to life-form evolution. The influence of the pathway of loading on other processes is discussed.     

On the role of the tricarboxylic acid cycle in plant productivity

The tricarboxylic acid(TCA) cycle is one of the canonical energy pathways of living systems, as well as being an example of a pathway in which dynamic enzyme assemblies, or metabolons, are well characterized. The role of the enzymes have been the subject of saturated transgenesis approaches, whereby the expression of the constituent enzymes were reduced or knocked out in order to ascertain their in vivo function.Some of the resultant plants exhibited improved photosynthesis and plant growth, under controlled greenhouse conditions. In addition, overexpression of the endogenous genes, or heterologous forms of a number of the enzymes, has been carried out in tomato fruit and the roots of a range of species, and in some instances improvement in fruit yield and postharvest properties and plant performance, under nutrient limitation, have been reported, respectively. Given a number of variants, in nature, we discuss possible synthetic approaches involving introducing these variants, or at least a subset of them, into plants. We additionally discuss the likely consequences of introducing synthetic metabolons, wherein certain pairs of reactions are artificially permanently assembled into plants, and speculate as to future strategies to further improve plant productivity by manipulation of the core metabolic pathway.     

Increasing modularity when downscaling networks from species to individuals

Downscaling networks from species to individuals is a useful approach to incorporate inter‐individual variation and to investigate whether topology of species‐based networks results from processes acting at the scale of individuals, such as foraging behaviour. Here, we analyzed pollen‐transport networks at two scales, i.e. pollinator species–plant species (sp–sp) and pollinator individuals–plant species (i–sp), and assessed whether modularity – a prevalent pattern in most pollination networks – is consistent across both scales. To test this we use three different algorithms developed for the calculation of modularity (unipartite, bipartite and weighted bipartite modularity) and compare the results obtained. Downscaling networks revealed a higher modular structure in i–sp networks than in sp–sp networks, regardless of the modular metric used. Using a null model approach, we show that modularity at the individual scale is originated by the existence of a high heterogeneity and specialization in the partition of pollen resources among conspecific individuals, a pattern which obviously cannot be observed at the species level. Modules in i–sp networks consisted of individuals sometimes neither taxonomically nor functionally related, but sharing common pollen resources at different moments of the flowering season. Interestingly, conspecific individuals may belong to different modules. Both plant and insect phenologies were important drivers of the modularity detected in individual‐based networks, even determining the topological roles of nodes in the networks. A temporal turnover of modules was identified, i.e. modules of individuals assembled and disassembled over time as species modify their foraging choices throughout the flowering season adjusting to ecological conditions. Downscaling from species to individual‐based networks is a promising approach to study the interplay among structural patterns and processes at different, but interdependent organizational levels.     

植物非叶绿色器官光合贡献研究进展

植物光合作用是生物界赖以生存的基础.长期以来,叶片被认为是植物进行光合作用的重要器官.然而在逆境条件下,植物非叶绿色器官的光合贡献也具有巨大的潜力.近年来,为了探究植物增产的新途径,科研工作者纷纷把目光投向了植物非叶绿色器官.本文简述了植物非叶绿色器官中能够进行光合作用的器官类型、非叶绿色器官光合贡献率及其光合贡献率的...     

Global spectroscopic analysis to study the regulation of the photosynthetic proton motive force: A critical reappraisal

In natural variable environments, plants rapidly adjust photosynthesis for optimum balance between photochemistry and photoprotection. These adjustments mainly occur via changes in their proton motive force (pmf). Recent studies based on time resolved analysis of the Electro Chromic Signal (ECS) bandshift of photosynthetic pigments in the model plant Arabidopsis thaliana have suggested an active role of ion fluxes across the thylakoid membranes in the regulation of the pmf. Among the different channels and transporters possibly involved in this phenomenon, we previously identified the TPK3 potassium channel. Plants silenced for TPK3 expression displayed light stress signatures, with reduced Non Photochemical Quenching (NPQ) capacity and sustained anthocyanin accumulation, even at moderate intensities. In this work we re-examined the role of this protein in pmf regulation, starting from the observation that both TPK3 knock-down (TPK3 KD) and WT plants display enhanced anthocyanin accumulation in the light under certain growth conditions, especially in old leaves. We thus compared the pmf features of young “green” (without anthocyanins) and old “red” (with anthocyanins) leaves in both genotypes using a global fit analysis of the ECS. We found that the differences in the ECS profile measured between the two genotypes reflect not only differences in TPK3 expression level, but also a modified photosynthetic activity of stressed red leaves, which are present in a larger amounts in the TPK3 KD plants.