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1.  Underwater Photosynthesis in Flooded Terrestrial Plants: A Matter of Leaf Plasticity 
Annals of Botany  2005;96(4):581-589.
• Background Flooding causes substantial stress for terrestrial plants, particularly if the floodwater completely submerges the shoot. The main problems during submergence are shortage of oxygen due to the slow diffusion rates of gases in water, and depletion of carbohydrates, which is the substrate for respiration. These two factors together lead to loss of biomass and eventually death of the submerged plants. Although conditions under water are unfavourable with respect to light and carbon dioxide supply, photosynthesis may provide both oxygen and carbohydrates, resulting in continuation of aerobic respiration.
• Scope This review focuses on evidence in the literature that photosynthesis contributes to survival of terrestrial plants during complete submergence. Furthermore, we discuss relevant morphological and physiological responses of the shoot of terrestrial plant species that enable the positive effects of light on underwater plant performance.
• Conclusions Light increases the survival of terrestrial plants under water, indicating that photosynthesis commonly occurs under these submerged conditions. Such underwater photosynthesis increases both internal oxygen concentrations and carbohydrate contents, compared with plants submerged in the dark, and thereby alleviates the adverse effects of flooding. Additionally, several terrestrial species show high plasticity with respect to their leaf development. In a number of species, leaf morphology changes in response to submergence, probably to facilitate underwater gas exchange. Such increased gas exchange may result in higher assimilation rates, and lower carbon dioxide compensation points under water, which is particularly important at the low carbon dioxide concentrations observed in the field. As a result of higher internal carbon dioxide concentrations in submergence-acclimated plants, underwater photorespiration rates are expected to be lower than in non-acclimated plants. Furthermore, the regulatory mechanisms that induce the switch from terrestrial to submergence-acclimated leaves may be controlled by the same pathways as described for heterophyllous aquatic plants.
PMCID: PMC4247027  PMID: 16024559
Flooding; gas exchange; heterophylly; hormonal regulation; leaf morphology; phenotypic plasticity; photorespiration; photosynthesis; Rumex; submergence; survival; wetlands
2.  A perspective on underwater photosynthesis in submerged terrestrial wetland plants 
AoB Plants  2011;2011:plr030.
Submergence inhibits photosynthesis by terrestrial wetland plants, but less so in species that possess leaf gas films when submerged. Floodwaters are often supersaturated with dissolved CO2 enabling photosynthesis by submerged terrestrial plants, although rates remain well-below those in air. This important adaptation that enhances survival in submerged conditions is reviewed.
Background and aims
Wetland plants inhabit flood-prone areas and therefore can experience episodes of complete submergence. Submergence impedes exchange of O2 and CO2 between leaves and the environment, and light availability is also reduced. The present review examines limitations to underwater net photosynthesis (PN) by terrestrial (i.e. usually emergent) wetland plants, as compared with submerged aquatic plants, with focus on leaf traits for enhanced CO2 acquisition.
Floodwaters are variable in dissolved O2, CO2, light and temperature, and these parameters influence underwater PN and the growth and survival of submerged plants. Aquatic species possess morphological and anatomical leaf traits that reduce diffusion limitations to CO2 uptake and thus aid PN under water. Many aquatic plants also have carbon-concentrating mechanisms to increase CO2 at Rubisco. Terrestrial wetland plants generally lack the numerous beneficial leaf traits possessed by aquatic plants, so submergence markedly reduces PN. Some terrestrial species, however, produce new leaves with a thinner cuticle and higher specific leaf area, whereas others have leaves with hydrophobic surfaces so that gas films are retained when submerged; both improve CO2 entry.
Submergence inhibits PN by terrestrial wetland plants, but less so in species that produce new leaves under water or in those with leaf gas films. Leaves with a thinner cuticle, or those with gas films, have improved gas diffusion with floodwaters, so that underwater PN is enhanced. Underwater PN provides sugars and O2 to submerged plants. Floodwaters often contain dissolved CO2 above levels in equilibrium with air, enabling at least some PN by terrestrial species when submerged, although rates remain well below those in air.
PMCID: PMC3249690  PMID: 22476500
3.  Biomechanical responses of aquatic plants to aerial conditions 
Annals of Botany  2013;112(9):1869-1878.
Background and Aims
Wetlands are impacted by changes in hydrological regimes that can lead to periods of low water levels. During these periods, aquatic plants experience a drastic change in the mechanical conditions that they encounter, from low gravitational and tensile hydrodynamic forces when exposed to flow under aquatic conditions, to high gravitational and bending forces under terrestrial conditions. The objective of this study was to test the capacity of aquatic plants to produce self-supporting growth forms when growing under aerial conditions by assessing their resistance to terrestrial mechanical conditions and the associated morpho-anatomical changes.
Plastic responses to aerial conditions were assessed by sampling Berula erecta, Hippuris vulgaris, Juncus articulatus, Lythrum salicaria, Mentha aquatica, Myosotis scorpioides, Nuphar lutea and Sparganium emersum under submerged and emergent conditions. The cross-sectional area and dry matter content (DMC) were measured in the plant organs that bear the mechanical forces, and their biomechanical properties in tension and bending were assessed.
Key Results
All of the species except for two had significantly higher stiffness in bending and thus an increased resistance to terrestrial mechanical conditions when growing under emergent conditions. This response was determined either by an increased allocation to strengthening tissues and thus a higher DMC, or by an increased cross-sectional area. These morpho-anatomical changes also resulted in increased strength and stiffness in tension.
The capacity of the studied species to colonize this fluctuating environment can be accounted for by a high degree of phenotypic plasticity in response to emersion. Further investigation is however needed to disentangle the finer mechanisms behind these responses (e.g. allometric relations, tissue make-up), their costs and adaptive value.
PMCID: PMC3838548  PMID: 24187030
Phenotypic plasticity; mechanical stress; biomechanics; stiffness; strength; aquatic herbaceous plants
4.  Molecular Adaptation of rbcL in the Heterophyllous Aquatic Plant Potamogeton 
PLoS ONE  2009;4(2):e4633.
Heterophyllous aquatic plants show marked phenotypic plasticity. They adapt to environmental changes by producing different leaf types: submerged, floating and terrestrial leaves. By contrast, homophyllous plants produce only submerged leaves and grow entirely underwater. Heterophylly and submerged homophylly evolved under selective pressure modifying the species-specific optima for photosynthesis, but little is known about the evolutionary outcome of habit. Recent evolutionary analyses suggested that rbcL, a chloroplast gene that encodes a catalytic subunit of RuBisCO, evolves under positive selection in most land plant lineages. To examine the adaptive evolutionary process linked to heterophylly or homophylly, we analyzed positive selection in the rbcL sequences of ecologically diverse aquatic plants, Japanese Potamogeton.
Principal Findings
Phylogenetic and maximum likelihood analyses of codon substitution models indicated that Potamogeton rbcL has evolved under positive Darwinian selection. The positive selection has operated specifically in heterophyllous lineages but not in homophyllous ones in the branch-site models. This suggests that the selective pressure on this chloroplast gene was higher for heterophyllous lineages than for homophyllous lineages. The replacement of 12 amino acids occurred at structurally important sites in the quaternary structure of RbcL, two of which (residue 225 and 281) were identified as potentially under positive selection.
Our analysis did not show an exact relationship between the amino acid replacements and heterophylly or homophylly but revealed that lineage-specific positive selection acted on the Potamogeton rbcL. The contrasting ecological conditions between heterophyllous and homophyllous plants have imposed different selective pressures on the photosynthetic system. The increased amino acid replacement in RbcL may reflect the continuous fine-tuning of RuBisCO under varying ecological conditions.
PMCID: PMC2646136  PMID: 19247501
5.  Blue light dose–responses of leaf photosynthesis, morphology, and chemical composition of Cucumis sativus grown under different combinations of red and blue light 
Journal of Experimental Botany  2010;61(11):3107-3117.
The blue part of the light spectrum has been associated with leaf characteristics which also develop under high irradiances. In this study blue light dose–response curves were made for the photosynthetic properties and related developmental characteristics of cucumber leaves that were grown at an equal irradiance under seven different combinations of red and blue light provided by light-emitting diodes. Only the leaves developed under red light alone (0% blue) displayed dysfunctional photosynthetic operation, characterized by a suboptimal and heterogeneously distributed dark-adapted Fv/Fm, a stomatal conductance unresponsive to irradiance, and a relatively low light-limited quantum yield for CO2 fixation. Only 7% blue light was sufficient to prevent any overt dysfunctional photosynthesis, which can be considered a qualitatively blue light effect. The photosynthetic capacity (Amax) was twice as high for leaves grown at 7% blue compared with 0% blue, and continued to increase with increasing blue percentage during growth measured up to 50% blue. At 100% blue, Amax was lower but photosynthetic functioning was normal. The increase in Amax with blue percentage (0–50%) was associated with an increase in leaf mass per unit leaf area (LMA), nitrogen (N) content per area, chlorophyll (Chl) content per area, and stomatal conductance. Above 15% blue, the parameters Amax, LMA, Chl content, photosynthetic N use efficiency, and the Chl:N ratio had a comparable relationship as reported for leaf responses to irradiance intensity. It is concluded that blue light during growth is qualitatively required for normal photosynthetic functioning and quantitatively mediates leaf responses resembling those to irradiance intensity.
PMCID: PMC2892149  PMID: 20504875
Blue light; chlorophyll fluorescence imaging; cucumber (Cucumis sativus); dose–response curves; leaf mass per unit leaf area (LMA); light-emitting diodes (LEDs); photoinhibition; photosynthetic capacity; red light; starch accumulation
6.  Hypersensitive to Red and Blue 1 and Its Modification by Protein Phosphatase 7 Are Implicated in the Control of Arabidopsis Stomatal Aperture 
PLoS Genetics  2012;8(5):e1002674.
The stomatal pores are located on the plant leaf epidermis and regulate CO2 uptake for photosynthesis and the loss of water by transpiration. Their stomatal aperture therefore affects photosynthesis, water use efficiency, and agricultural crop yields. Blue light, one of the environmental signals that regulates the plant stomatal aperture, is perceived by the blue/UV-A light-absorbing cryptochromes and phototropins. The signal transduction cascades that link the perception of light to the stomatal opening response are still largely unknown. Here, we report two new players, Hypersensitive to Red and Blue 1 (HRB1) and Protein Phosphatase 7 (PP7), and their genetic and biochemical interactions in the control of stomatal aperture. Mutations in either HRB1 or PP7 lead to the misregulation of the stomatal aperture and reduce water loss under blue light. Both HRB1 and PP7 are expressed in the guard cells in response to a light-to-dark or dark-to-light transition. HRB1 interacts with PP7 through its N-terminal ZZ-type zinc finger motif and requires a functional PP7 for its stomatal opening response. HRB1 is phosphorylated in vivo, and PP7 can dephosphorylate HRB1. HRB1 is mostly dephosphorylated in a protein complex of 193 kDa in the dark, and blue light increases complex size to 285 kDa. In the pp7 mutant, this size shift is impaired, and HRB1 is predominately phosphorylated. We propose that a modification of HRB1 by PP7 under blue light is essential to acquire a proper conformation or to bring in new components for the assembly of a functional HRB1 protein complex. Guard cells control stomatal opening in response to multiple environmental or biotic stimuli. This study may furnish strategies that allow plants to enjoy the advantages of both constitutive and ABA-induced protection under water-limiting conditions.
Author Summary
Stomatal aperture is regulated by many environmental and biotic cues such as blue light, drought, elevated CO2 concentrations, high humidity, and pathogenic elicitors. Stomatal apertures vary over diurnal cycles, and stomata tend to be open during the day in response to blue light and tend to be closed at night. The blue/UV-A light-absorbing cryptochromes and phototropins are the receptors for the blue light response. We report the action of HRB1, a nuclear ZZ-type zinc finger protein, and PP7, a positive regulator of blue light signaling in the nucleus, in the signal transduction cascades downstream of blue light perception. Both hrb1 and pp7 mutants are more resistant to dehydration and show reductions in both water loss and blue light-regulated stomatal aperture. Our studies on their genetic and biochemical interactions offer novel insights on the network structure of the light signaling machinery and plant interactions with the environment. Periodic drought is one of the major environmental factors that limits biomass production and crop yield in a changing global climate. Our studies may open new possibilities to engineer plants to survive desiccation.
PMCID: PMC3349726  PMID: 22589732
7.  Co-ordination of hydraulic and stomatal conductances across light qualities in cucumber leaves 
Journal of Experimental Botany  2011;63(3):1135-1143.
Long-term effects of light quality on leaf hydraulic conductance (Kleaf) and stomatal conductance (gs) were studied in cucumber, and their joint impact on leaf photosynthesis in response to osmotic-induced water stress was assessed. Plants were grown under low intensity monochromatic red (R, 640 nm), blue (B, 420 nm) or combined red and blue (R:B, 70:30) light. Kleaf and gs were much lower in leaves that developed without blue light. Differences in gs were caused by differences in stomatal aperture and stomatal density, of which the latter was largely due to differences in epidermal cell size and hardly due to stomatal development. Net photosynthesis (AN) was lowest in R-, intermediate in B-, and highest in RB- grown leaves. The low AN in R-grown leaves correlated with a low leaf internal CO2 concentration and reduced PSII operating efficiency. In response to osmotic stress, all leaves showed similar degrees of stomatal closure, but the reduction in AN was larger in R- than in B- and RB-grown leaves. This was probably due to damage of the photosynthetic apparatus, which only occurred in R-grown leaves. The present study shows the co-ordination of Kleaf and gs across different light qualities, while the presence of blue in the light spectrum seems to drive both Kleaf and gs towards high, sun-type leaf values, as was previously reported for maximal photosynthetic capacity and leaf morphology. The present results suggest the involvement of blue light receptors in the usually harmonized development of leaf characteristics related to water relations and photosynthesis under different light environments.
PMCID: PMC3276089  PMID: 22121201
Amphistomatous; Cucumis sativus; leaf development; leaf hydraulic conductance; light quality; osmotic stress; photosynthesis; stomatal conductance; stomatal density; stomatal opening
8.  Association between winter anthocyanin production and drought stress in angiosperm evergreen species 
Journal of Experimental Botany  2010;61(6):1699-1709.
Leaves of many evergreen angiosperm species turn red under high light during winter due to the production of anthocyanin pigments, while leaves of other species remain green. There is currently no explanation for why some evergreen species exhibit winter reddening while others do not. Conditions associated with low leaf water potentials (Ψ) have been shown to induce reddening in many plant species. Because evergreen species differ in susceptibility to water stress during winter, it is hypothesized that species which undergo winter colour change correspond with those that experience/tolerate the most severe daily declines in leaf Ψ during winter. Six angiosperm evergreen species which synthesize anthocyanin in leaves under high light during winter and five species which do not were studied. Field Ψ, pressure/volume curves, and gas exchange measurements were derived in summer (before leaf colour change had occurred) and winter. Consistent with the hypothesis, red-leafed species as a group had significantly lower midday Ψ in winter than green-leafed species, but not during the summer when all the leaves were green. However, some red-leafed species showed midday declines similar to those of green-leafed species, suggesting that low Ψ alone may not induce reddening. Pressure–volume curves also provided some evidence of acclimation to more negative water potentials by red-leafed species during winter (e.g. greater osmotic adjustment and cell wall hardening on average). However, much overlap in these physiological parameters was observed as well between red and green-leafed species, and some of the least drought-acclimated species were red-leafed. No difference was observed in transpiration (E) during winter between red and green-leaved species. When data were combined, only three of the six red-leafed species examined appeared physiologically acclimated to prolonged drought stress, compared to one of the five green-leafed species. This suggests that drought stress alone is not sufficient to explain winter reddening in evergreen angiosperms.
PMCID: PMC2852665  PMID: 20202995
Anthocyanin; carbohydrates; drought; evergreen; osmolarity; photosynthesis; sugars; winter; water relations
9.  Influence of Green, Red and Blue Light Emitting Diodes on Multiprotein Complex Proteins and Photosynthetic Activity under Different Light Intensities in Lettuce Leaves (Lactuca sativa L.) 
The objective of this study was to investigate the response of light emitting diodes (LEDs) at different light intensities (70 and 80 for green LEDs, 88 and 238 for red LEDs and 80 and 238 μmol m−2 s−1 for blue LEDs) at three wavelengths in lettuce leaves. Lettuce leaves were exposed to (522 nm), red (639 nm) and blue (470 nm) LEDs of different light intensities. Thylakoid multiprotein complex proteins and photosynthetic metabolism were then investigated. Biomass and photosynthetic parameters increased with an increasing light intensity under blue LED illumination and decreased when illuminated with red and green LEDs with decreased light intensity. The expression of multiprotein complex proteins including PSII-core dimer and PSII-core monomer using blue LEDs illumination was higher at higher light intensity (238 μmol m−2 s−1) and was lowered with decreased light intensity (70–80 μmol m−2 s−1). The responses of chloroplast sub-compartment proteins, including those active in stomatal opening and closing, and leaf physiological responses at different light intensities, indicated induced growth enhancement upon illumination with blue LEDs. High intensity blue LEDs promote plant growth by controlling the integrity of chloroplast proteins that optimize photosynthetic performance in the natural environment.
PMCID: PMC3975419  PMID: 24642884
lettuce (Lactuca sativa L.); light emitting diodes; photosynthesis; stomata; BN-PAGE
10.  Photosynthetic acclimation is important for post-submergence recovery of photosynthesis and growth in two riparian species 
Annals of Botany  2009;104(7):1435-1444.
Background and Aims
Concomitant increases in O2 and irradiance upon de-submergence can cause photoinhibition and photo-oxidative damage to the photosynthetic apparatus of plants. As energy and carbohydrate supply from photosynthesis is needed for growth, it was hypothesized that post-submergence growth recovery may require efficient photosynthetic acclimation to increased O2 and irradiance to minimize photo-oxidative damage. The hypothesis was tested in two flood-tolerant species: a C3 herb, Alternanthera philoxeroides; and a C4 grass, Hemarthria altissima. The impact of low O2 and low light, typical conditions in turbid floodwater, on post-submergence recovery was assessed by different flooding treatments combined with shading.
Experiments were conducted during 30 d of flooding (waterlogging or submergence) with or without shading and subsequent recovery of 20 d under growth conditions. Changes in dry mass, number of branches/tillers, and length of the longest internodes and main stems were recorded to characterize growth responses. Photosynthetic parameters (photosystem II efficiency and non-photochemical quenching) were determined in mature leaves based on chlorophyll a fluorescence measurements.
Key Results
In both species growth and photosynthesis recovered after the end of the submergence treatment, with recovery of photosynthesis (starting shortly after de-submergence) preceding recovery of growth (pronounced on days 40–50). The effective quantum yield of photosystem II and non-photochemical quenching were diminished during submergence but rapidly increased upon de-submergence. Similar changes were found in all shaded plants, with or without flooding. Submerged plants did not suffer from photoinhibition throughout the recovery period although their growth recovery was retarded.
After sudden de-submergence the C3 plant A. philoxeroides and the C4 plant H. altissima were both able to maintain the functionality of the photosynthetic apparatus through rapid acclimation to changing O2 and light conditions. The ability for photosynthetic acclimation may be essential for adaptation to wetland habitats in which water levels fluctuate.
PMCID: PMC2778401  PMID: 19854720
Aerenchyma; Alternanthera philoxeroides; flooding; growth; Hemarthria altissima; low light; photosynthesis; shade; submergence; waterlogging; wetland plant
11.  Light controls phospholipase A2α and β gene expression in Citrus sinensis 
Journal of Experimental Botany  2010;61(9):2469-2478.
The low-molecular weight secretory phospholipase A2α (CssPLA2α) and β (CsPLA2β) cloned in this study exhibited diurnal rhythmicity in leaf tissue of Citrus sinensis. Only CssPLA2α displayed distinct diurnal patterns in fruit tissues. CssPLA2α and CsPLA2β diurnal expression exhibited periods of approximately 24 h; CssPLA2α amplitude averaged 990-fold in the leaf blades from field-grown trees, whereas CsPLA2β amplitude averaged 6.4-fold. Diurnal oscillation of CssPLA2α and CsPLA2β gene expression in the growth chamber experiments was markedly dampened 24 h after transfer to continuous light or dark conditions. CssPLA2α and CsPLA2β expressions were redundantly mediated by blue, green, red and red/far-red light, but blue light was a major factor affecting CssPLA2α and CsPLA2β expression. Total and low molecular weight CsPLA2 enzyme activity closely followed diurnal changes in CssPLA2α transcript expression in leaf blades of seedlings treated with low intensity blue light (24 μmol m−2 s−1). Compared with CssPLA2α basal expression, CsPLA2β expression was at least 10-fold higher. Diurnal fluctuation and light regulation of PLA2 gene expression and enzyme activity in citrus leaf and fruit tissues suggests that accompanying diurnal changes in lipophilic second messengers participate in the regulation of physiological processes associated with phospholipase A2 action.
PMCID: PMC2877900  PMID: 20388744
Citrus sinensis; diurnal; light regulation; phospholipase A
12.  Cessation of Tillering in Spring Wheat in Relation to Light Interception and Red : Far-red Ratio 
Annals of Botany  2006;97(4):649-658.
• Background and Aims The production of axillary shoots (tillering) in spring wheat (Triticum aestivum) depends on intraspecific competition. The mechanisms that underlie this competition are complex, but light within the wheat canopy plays a key role. The main objectives of this paper are to analyse the effects of plant population density and shade on tillering dynamics of spring wheat, to assess the canopy conditions quantitatively at the time of tillering cessation, and to analyse the relationship between the tiller bud and the leaf on the same phytomer.
• Methods Spring wheat plants were grown at three plant population densities and under two light regimes (25 % and 100 % light). Tiller appearance, fraction of the light intercepted, and red : far-red ratio at soil level were recorded. On six sampling dates the growth status of axillary buds was analysed.
• Key Results Tillering ceased earlier at high population densities and ceased earlier in the shade than in full sunlight. At cessation of tillering, both the fraction of light intercepted and the red : far-red ratio at soil level were similar in all treatments. Leaves on the same phytomer of buds that grew out showed more leaf mass per unit area than those on the same phytomer of buds that remained dormant.
• Conclusions Tillering ceases at specific light conditions within the wheat canopy, independent of population density, and to a lesser extent independent of light intensity. It is suggested that cessation of tillering is induced when the fraction of PAR intercepted by the canopy exceeds a specific threshold (0·40–0·45) and red : far-red ratio drops below 0·35–0·40.
PMCID: PMC2803660  PMID: 16464875
Triticum aestivum; wheat; tiller; bud; plant population density; shade; PAR; red : far-red ratio; functional–structural model
13.  A Dynamic Analysis of the Shade-induced Plasticity in Arabidopsis thaliana Rosette Leaf Development Reveals New Components of the Shade-adaptative Response 
Annals of Botany  2006;97(3):443-452.
• Background and Aims It is well known that plant aerial development is affected by light intensity in terms of the date of flowering, the length of stems and petioles, and the final individual leaf area. The aim of the work presented here was to analyse how shade-induced changes in leaf development occur on a dynamic basis from the whole rosette level to that of the cells.
• Methods Care was taken to ensure that light intensity was the only source of micro-meteorological variation in the study. The dynamics of leaf production, rosette expansion, individual leaf area expansion and epidermal cell expansion were analysed in Arabidopsis thaliana plants grown under two light intensities in three independent experiments.
• Key Results The total area of rosette leaves was reduced by the shading treatment. Both the number of leaves produced and their individual leaf areas were reduced. The reduction in leaf number was associated with a reduction in leaf initiation rate and the duration of the phase of leaf production. The reduction in individual leaf area was associated with a reduction in leaf expansion rate and an increase in the duration of leaf expansion. The changes in leaf expansion dynamics were accompanied by a decrease in epidermal cell number which was partly compensated for by an increase in epidermal cell area. Overall, the whole rosette leaf expansion rate was reduced by shading, whereas the total duration of rosette leaf expansion was unaffected. This was mainly due to the accumulation of the increases in the durations of expansion of each individual leaf which was associated with an increase in cell expansion.
• Conclusions The dynamic analysis presented here reveals a new shade-adaptative response mediated via the control of area expansion at the cell, organ and whole plant levels.
PMCID: PMC2803649  PMID: 16371443
Arabidopsis thaliana; light intensity; leaf development; expansion rate; duration; initiation; cell expansion
14.  Multi-level Modeling of Light-Induced Stomatal Opening Offers New Insights into Its Regulation by Drought 
PLoS Computational Biology  2014;10(11):e1003930.
Plant guard cells gate CO2 uptake and transpirational water loss through stomatal pores. As a result of decades of experimental investigation, there is an abundance of information on the involvement of specific proteins and secondary messengers in the regulation of stomatal movements and on the pairwise relationships between guard cell components. We constructed a multi-level dynamic model of guard cell signal transduction during light-induced stomatal opening and of the effect of the plant hormone abscisic acid (ABA) on this process. The model integrates into a coherent network the direct and indirect biological evidence regarding the regulation of seventy components implicated in stomatal opening. Analysis of this signal transduction network identified robust cross-talk between blue light and ABA, in which [Ca2+]c plays a key role, and indicated an absence of cross-talk between red light and ABA. The dynamic model captured more than 1031 distinct states for the system and yielded outcomes that were in qualitative agreement with a wide variety of previous experimental results. We obtained novel model predictions by simulating single component knockout phenotypes. We found that under white light or blue light, over 60%, and under red light, over 90% of all simulated knockouts had similar opening responses as wild type, showing that the system is robust against single node loss. The model revealed an open question concerning the effect of ABA on red light-induced stomatal opening. We experimentally showed that ABA is able to inhibit red light-induced stomatal opening, and our model offers possible hypotheses for the underlying mechanism, which point to potential future experiments. Our modelling methodology combines simplicity and flexibility with dynamic richness, making it well suited for a wide class of biological regulatory systems.
Author Summary
Stomata are microscopic pores surrounded and regulated by pairs of guard cells located on the surface of plant leaves. Stomata participate in CO2 uptake, O2 release and water vapor loss. Blue and red light induce stomatal opening (enlargement of the pores), which allows the uptake of CO2, providing the raw material for photosynthesis, and the release of O2 into the atmosphere. The stress hormone abscisic acid induces minimization of pore width, decreasing water loss through transpiration. During drought conditions these counteracting stimuli jointly determine the overall stomatal movement through an integrated guard cell signalling cascade. We synthesized this interaction network between blue light, red light, and abscisic acid by aggregating and interpreting the abundant biological evidence that has been accumulated to date. We used the resulting network as the basis of a multi-level dynamic model of stomatal opening regulation in response to multiple stimuli. The model is validated by comparing its results to a large number of published experimental observations. Our model, and our experiments inspired by it, reveal an unexplored facet of the interplay between light and abscisic acid in guard cell signalling. The model directs future experiments, and its methodology can readily be applied to other systems.
PMCID: PMC4230748  PMID: 25393147
15.  Submerged in darkness: adaptations to prolonged submergence by woody species of the Amazonian floodplains 
Annals of Botany  2008;103(2):359-376.
In Amazonian floodplain forests, >1000 tree species grow in an environment subject to extended annual submergence which can last up to 9 months each year. Water depth can reach 10 m, fully submerging young and also adult trees, most of which reproduce during the flood season. Complete submergence occurs regularly at the seedling or sapling stage for many species that colonize low-lying positions in the flooding gradient. Here hypoxic conditions prevail close to the water surface in moving water, while anaerobic conditions are common in stagnant pools. Light intensities in the floodwater are very low.
Questions and Aims
Despite a lack of both oxygen and light imposed by submergence for several months, most leafed seedlings survive. Furthermore, underwater growth has also been observed in several species in the field and under experimental conditions. The present article assesses how these remarkable plants react to submergence and discusses physiological mechanisms and anatomical adaptations that may explain their success.
PMCID: PMC2707320  PMID: 19001429
Adaptation; Amazonian floodplains; darkness; environmental stress; flooding; hypoxia; submergence tolerance; trees; underwater photosynthesis; woody species
16.  Investigations on the Photoregulation of Chloroplast Movement and Leaf Positioning in Arabidopsis 
Plant and Cell Physiology  2012;54(1):48-56.
We recently investigated the roles of the phototropin 1 (PHOT1) LOV (light, oxygen or voltage) domains in mediating phototropic curvature in transgenic Arabidopsis seedlings expressing either wild-type PHOT1 or PHOT1 with one or both LOV domains inactivated by a single amino acid replacement. We have now investigated the role of the PHOT1 LOV domains in chloroplast movement and in leaf positioning in response to blue light. Low fluence rate blue light is known to mediate a chloroplast accumulation response and high fluence rate blue light an avoidance response in Arabidopsis leaves. As was the case for phototropism, LOV2 of PHOT1 is essential for chloroplast accumulation and LOV1 is dispensable. PHOT1 LOV2 is also essential to maintain developing primary leaves in a horizontal position under white light from above and LOV1 is again dispensable. A red light pulse given to dark-adapted light-grown plants followed by 2 h of darkness enhances both the chloroplast accumulation response under dim blue light and the chloroplast avoidance response under strong blue light. The effect is far-red reversible. This photoreversible response is normal in a phyB null mutant but does not appear in a phyA null mutant. These results suggest that phyA mediates the enhancement, induced by a red light pulse, of blue light-induced chloroplast movements.
PMCID: PMC3539441  PMID: 22782888
Arabidopsis; Chloroplast; movement; LOV; domain; Phototropin; Phytochrome
17.  Physiological analyses of traits associated with tolerance of long-term partial submergence in rice 
AoB Plants  2014;6:plu058.
Long-term stagnant flooding (SF, 50 cm water depth) greatly reduces rice yield. We assessed physiological mechanisms associated with SF tolerance in contrasting rice genotypes. SF reduced yield by 47% due to low biomass caused by reduced light interception and leaf growth above water; and reduced panicle number by 52% because of low tillering. Shoot elongation correlated positively with leaf growth and biomass production, but negatively with stem nonstructural carbohydrates (NSC). Tolerant varieties were either inherently tall or elongate moderately (<2.0 cm d−1) with rising floodwater. Optimum shoot elongation with rising floodwater is therefore a priority for future breeding work.
Floods are major constraints to crop production worldwide. In low-lying, flood-prone areas of the tropics, longer-term partial submergence (stagnant flooding [SF]) greatly reduces rice yield. This study assesses shoot growth and several physiological mechanisms associated with SF tolerance in rice. Five rice genotypes with contrasting responses to SF were evaluated in field ponds. Following transplanting, floodwater was gradually increased at a rate of ∼2 cm day−1 to reach a final depth of 50 cm and then maintained until maturity. Although plants were not fully submerged, the yield was reduced by 47 % across genotypes compared with those grown under control conditions (6.1 vs. 3.3 t ha−1). This reduction was mainly attributed to the reduction in biomass caused by reduced light interception and leaf growth above the water. Stagnant flooding also reduced panicle number per unit area by 52 % because of reduced tillering. Shoot elongation rate kept pace with rising floodwater and correlated positively with leaf growth and biomass production. Conversely, stem non-structural carbohydrate (NSC) concentration correlated negatively with shoot elongation rate, suggesting that fast-elongating genotypes actively consume NSCs to avoid complete submergence. Moderate shoot elongation rate strongly and positively correlated with grain yield under SF; however, elongation at rates >2.0 cm day−1 was associated with reduced harvest index due to a smaller panicle size and increased lodging. Tolerant varieties were found to be either inherently tall or elongate moderately with rising floodwater. Our studies suggest that to improve tolerance of SF an appropriate phenotype should combine both of these traits. Fine-tuning for optimum shoot elongation with rising floodwater is, therefore, a priority for future work.
PMCID: PMC4216431  PMID: 25270231
Flooding stress; rainfed lowland rice; shoot elongation; stagnant flooding; tillering
18.  Use of Confocal Laser as Light Source Reveals Stomata-Autonomous Function 
PLoS ONE  2006;1(1):e36.
In most terrestrial plants, stomata open during the day to maximize the update of CO2 for photosynthesis, but they close at night to minimize water loss. Blue light, among several environmental factors, controls this process. Stomata response to diverse stimuli seems to be dictated by the behaviour of neighbour stomata creating leaf areas of coordinated response. Here individual stomata of Arabidopsis leaves were illuminated with a short blue-light pulse by focusing a confocal argon laser. Beautifully, the illuminated stomata open their pores, whereas their dark-adapted neighbours unexpectedly experience no change. This induction of individual stomata opening by low fluence rates of blue light was disrupted in the phototropin1 phototropin2 (phot1 phot2) double mutant, which exhibits insensitivity of stomatal movements in blue-illuminated epidermal strips. The irradiation of all epidermal cells making direct contact with a given stoma in both wild type and phot1 phot2 plants does not trigger its movement. These results unravel the stoma autonomous function in the blue light response and illuminate the implication of PHOT1 and/or PHOT2 in such response. The micro spatial heterogeneity that solar blue light suffers in partially shaded leaves under natural conditions highlights the physiological significance of the autonomous stomatal behaviour.
PMCID: PMC1762337  PMID: 17183664
19.  Effect of light on the growth and photosynthesis of an invasive shrub in its native range 
AoB Plants  2014;6:plu033.
We studied the growth and photosynthetic capacity of Berberis darwinii shrub growing under different light conditions in their native area of Argentina to test if plant physiology differs from invaded area (using studies carried out in New Zealand). In its native range B. darwinii grows under different light conditions by adjusting shoot and leaf morphology and physiology. Plants of B. darwinii growing under the same light environments show similar physiology in native and invasive ranges. Therefore, intra-specific variations of the functional traits in native area do not condition successful invasiveness.
Invasive species' success may depend on ecophysiological attributes present in their native area or those derived from changes that took place in the invaded environment. We studied the growth and photosynthetic capacity of Berberis darwinii shrubs growing under different light conditions (gap, forest edge and below the canopy) in their native area of Patagonia, Argentina. Leaf photosynthesis results determined in the native area were discussed in relation to information provided by studies carried out under the same light conditions in an invaded area in New Zealand. Shoot elongation, leaf production, stem and leaf biomass per shoot, and specific leaf area (SLA, cm2 g−1) were determined in five adult plants, randomly selected in each of the three light conditions at two forest sites. Net photosynthesis as a function of PPFD (photosynthetic photon flux density), stomatal conductance (gs), maximum light-saturated photosynthesis rate (Pmax), Pmass (on mass bases) and water-use efficiency (WUEi) were determined in plants of one site. We predicted that functional traits would differ between populations of native and invasive ranges. In their native area, plants growing under the canopy produced the longest shoots and had the lowest values for shoot emergence and foliar biomass per shoot, while their SLA was higher than gap and forest edge plants. Leaf number and stem biomass per shoot were independent of light differences. Leaves of gap plants showed higher Pmax, Pmass and gs but lower WUEi than plants growing at the forest edge. In its native range B. darwinii grows under different light conditions by adjusting shoot and leaf morphology and physiology. Plants of B. darwinii growing under the same light conditions show similar physiology in native and invasive ranges. This means that for B. darwinii, intra-specific variation of the functional traits studied here does not condition successful spread in new areas.
PMCID: PMC4240251  PMID: 24969502
Berberis darwinii; ecophysiological attributes; light environments; native and invasion area; plant invasion.
20.  Diel leaf growth of soybean: a novel method to analyze two-dimensional leaf expansion in high temporal resolution based on a marker tracking approach (Martrack Leaf) 
Plant Methods  2013;9:30.
We present a novel method for quantitative analysis of dicot leaf expansion at high temporal resolution. Image sequences of growing leaves were assessed using a marker tracking algorithm. An important feature of the method is the attachment of dark beads that serve as artificial landmarks to the leaf margin. The beads are mechanically constricted to the focal plane of a camera. Leaf expansion is approximated by the increase in area of the polygon defined by the centers of mass of the beads surrounding the leaf. Fluctuating illumination conditions often pose serious problems for tracking natural structures of a leaf; this problem is circumvented here by the use of the beads.
The new method has been used to assess leaf growth in environmental situations with different illumination conditions that are typical in agricultural and biological experiments: Constant illumination via fluorescent light tubes in a climate chamber, a mix of natural and artificial illumination in a greenhouse and natural illumination of the situation on typical summer days in the field. Typical features of diel (24h) soybean leaf growth patterns were revealed in all three conditions, thereby demonstrating the general applicability of the method. Algorithms are provided to the entire community interested in using such approaches.
The implementation Martrack Leaf presented here is a robust method to investigate diel leaf growth rhythms both under natural and artificial illumination conditions. It will be beneficial for the further elucidation of genotype x environment x management interactions affecting leaf growth processes.
PMCID: PMC3750653  PMID: 23883317
Marker tracking; Phenotyping; Image analysis; Plant growth; Diel growth; Natural illumination
21.  Bud Composition, Branching Patterns and Leaf Phenology in Cerrado Woody Species 
Annals of Botany  2005;96(6):1075-1084.
• Background and Aims Plants have complex mechanisms of aerial biomass exposition, which depend on bud composition, the period of the year in which shoot extension occurs, branching pattern, foliage persistence, herbivory and environmental conditions.
• Methods The influence of water availability and temperature on shoot growth, the bud composition, the leaf phenology, and the relationship between partial leaf fall and branching were evaluated over 3 years in Cerrado woody species Bauhinia rufa (BR), Leandra lacunosa (LL) and Miconia albicans (MA).
• Key Results Deciduous BR preformed organs in buds and leaves flush synchronously at the transition from the dry to the wet season. The expansion time of leaves is <1 month. Main shoots (first-order axis, A1 shoots) extended over 30 d and they did not branch. BR budding and foliage unfolds were brought about independently of inter-annual rainfall variations. By contrast, in LL and MA evergreen species, the shoot extension rate and the neoformation of aerial organs depended on rainfall. Leaf emergence was continuous for 2–6 months and lamina expansion took place over 1–4 months. The leaf life span was 5–20 months and the main A1 shoot extension happened over 122–177 d. Both evergreen species allocated biomass to shoots, leaves or flowers continuously during the year, branching in the middle of the wet season to form second-order (A2 shoots) and third-order (A3 shoots) axis in LL and A2 shoots in MA. Partial shed of A1 shoot leaves would facilitate a higher branching intensity A2 shoot production in LL than in MA. MA presented a longer leaf life span, produced a lower percentage of A2 shoots but had a higher meristem persistence on A1 and A2 shoots than LL.
• Conclusions It was possible to identify different patterns of aerial growth in Cerrado woody species defined by shoot-linked traits such as branching pattern, bud composition, meristem persistence and leaf phenology. These related traits must be considered over and above leaf deciduousness for searching functional guilds in a Cerrado woody community. For the first time a relationship between bud composition, shoot growth and leaf production pattern is found in savanna woody plants.
PMCID: PMC4247095  PMID: 16157631
Bauhinia rufa; branching; Brazil; bud composition; Cerrado; flowering; leaf phenology; Leandra lacunosa; meristem persistence; Miconia albicans; synchronic leaves production; continuous leaf production
22.  Acclimatory responses of Arabidopsis to fluctuating light environment: comparison of different sunfleck regimes and accessions 
Photosynthesis Research  2012;113(1-3):221-237.
Acclimation to fluctuating light environment with short (lasting 20 s, at 650 or 1,250 μmol photons m−2 s−1, every 6 or 12 min) or long (for 40 min at 650 μmol photons m−2 s−1, once a day at midday) sunflecks was studied in Arabidopsis thaliana. The sunfleck treatments were applied in the background daytime light intensity of 50 μmol photons m−2 s−1. In order to distinguish the effects of sunflecks from those of increased daily irradiance, constant light treatments at 85 and 120 μmol photons m−2 s−1, which gave the same photosynthetically active radiation (PAR) per day as the different sunfleck treatments, were also included in the experiments. The increased daily total PAR in the two higher constant light treatments enhanced photosystem II electron transport and starch accumulation in mature leaves and promoted expansion of young leaves in Columbia-0 plants during the 7-day treatments. Compared to the plants remaining under 50 μmol photons m−2 s−1, application of long sunflecks caused upregulation of electron transport without affecting carbon gain in the form of starch accumulation and leaf growth or the capacity of non-photochemical quenching (NPQ). Mature leaves showed marked enhancement of the NPQ capacity under the conditions with short sunflecks, which preceded recovery and upregulation of electron transport, demonstrating the initial priority of photoprotection. The distinct acclimatory responses to constant PAR, long sunflecks, and different combinations of short sunflecks are consistent with acclimatory adjustment of the processes in photoprotection and carbon gain, depending on the duration, frequency, and intensity of light fluctuations. While the responses of leaf expansion to short sunflecks differed among the seven Arabidopsis accessions examined, all plants showed NPQ upregulation, suggesting limited ability of this species to utilize short sunflecks. The increase in the NPQ capacity was accompanied by reduced chlorophyll contents, higher levels of the xanthophyll-cycle pigments, faster light-induced de-epoxidation of violaxanthin to zeaxanthin and antheraxanthin, increased amounts of PsbS protein, as well as enhanced activity of superoxide dismutase. These acclimatory mechanisms, involving reorganization of pigment–protein complexes and upregulation of other photoprotective reactions, are probably essential for Arabidopsis plants to cope with photo-oxidative stress induced by short sunflecks without suffering from severe photoinhibition and lipid peroxidation.
PMCID: PMC3430843  PMID: 22729524
Acclimation; Arabidopsis thaliana; Non-photochemical quenching; Photoprotection; Sunflecks; Xanthophyll cycle
23.  Phenotypic variation in heteroblastic woody species does not contribute to shade survival 
AoB Plants  2011;2011:plr013.
Heteroblastic species change their leaf morphology due to changes in light environment. However, growth and biomass allocation pattern do not contribute to their better survival relative to homoblastic congeners in low light. Thus, shade does not select for leaf heteroblasty.
Background and aims
Leaf heteroblasty involves dramatic phenotypic differences between adult and seedling leaves while leaves of homoblastic plants display only small differences. This study tested whether, in low-light environments, the marked difference in the morphology of seedling leaves that characterizes heteroblastic species confers advantages for seedling survival and growth compared with homoblastic congeners.
Four pairs of heteroblastic and homoblastic species in genera Hoheria, Aristotelia, Pseudopanax and Melicope were grown in simulated full sunlight (100 % of light, red:far red ratio (R:FR) = 1.25) or in simulated forest understorey shade (5 % of full sunlight, R:FR ratio = 0.25) in a glasshouse.
Principal results
After 9 months, 100 % of seedlings of both homoblastic and heteroblastic species survived in full sun while in the understorey treatment there were 25 % fewer heteroblastic survivors than homoblastic congeners. Compared with homoblastic congeners, all heteroblastic species except for Pseudopanax crassifolius produced more and smaller leaves and branches, but grew more slowly in height, root collar diameter and total biomass both in full sun and in forest understorey treatments.
Homoblastic species survive and grow better in the forest understorey light treatment, suggesting that heteroblastic seedling leaf morphology does not give an advantage over homoblastic congeners under low light intensities.
PMCID: PMC3129537  PMID: 22476483
24.  Rapid and Pervasive Occupation of Fallen Mangrove Leaves by a Marine Zoosporic Fungus † 
Applied and Environmental Microbiology  1987;53(10):2464-2469.
Samples of leaves of red mangrove (Rhizophora mangle) were incubated on an agar medium selective for pythiaceous oomycetes. Leaves on trees above the water did not contain oomycetes. Marine oomycetes, principally Phytophthora vesicula, had colonized leaves within 2 h of leaf submergence, probably finding them by chemotaxis. The frequency of occurrence of P. vesicula in submerged leaves reached 100% within 30 h of submergence. By 43 h most, if not all, parts of leaves were occupied, and surface treatment with a biocide indicated that leaves were occupied internally. Frequencies of P. vesicula remained near 100% through about 2 weeks of submergence and then declined to about 60% in older (≥4 weeks) leaves. Leaves of white mangrove (Laguncularia racemosa) were also extensively occupied by P. vesicula after falling into the water column, but decaying leaves of turtlegrass (Thalassia testudinum) were not colonized by oomycetes. Ergosterol analysis indicated that the standing crop of living, non-oomycete (ergosterol-containing) fungal mass in submerged red-mangrove leaves did not rise above that which had been present in senescent leaves on the tree; decaying turtlegrass leaves had an ergosterol content that was only about 2% of the maximum concentration detected for red-mangrove leaves. These results suggest that oomycetes are the predominant mycelial eucaryotic saprotrophs of mangrove leaves that fall into the water column and that for turtlegrass leaves which live, die, and decompose under submerged conditions, mycelial eucaryotes make no substantial contribution to decomposition.
PMCID: PMC204130  PMID: 16347463
25.  The core interthreshold zone during exposure to red and blue light 
This study tested the hypothesis that the core interthreshold zone (CIZ) changes during exposure to red or blue light via the non-visual pathway, because it is known that light intensity affects the central nervous system. We conducted a series of human experiments with 5 or 10 male subjects in each experiment.
The air temperature in the climatic chamber was maintained at 20 to 24°C. The subjects wore suits perfused with 25°C water at a rate of 600 cm3/min. They exercised on an ergometer at 50% of their maximum work rate for 10 to 15 minutes until sweating commenced, and then remained continuously seated without exercise until their oxygen uptake increased. The rectal temperature and skin temperatures at four sites were monitored using thermistors. The sweating rate was measured at the forehead with a sweat rate monitor. Oxygen uptake was monitored with a gas analyzer. The subjects were exposed to red or blue light at 500 lx and 1000 lx in both summer and winter.
The mean CIZs at 500 lx were 0.23 ± 0.16°C under red light and 0.20 ± 0.10°C under blue light in the summer, and 0.19 ± 0.20°C under red light and 0.26 ± 0.24°C under blue light in the winter. The CIZs at 1000 lx were 0.18 ± 0.14°C under red light and 0.15 ± 0.20°C under blue light in the summer, and 0.52 ± 0.18°C under red light and 0.71 ± 0.28°C under blue light in the winter. A significant difference (P <0.05) was observed in the CIZs between red and blue light at 1000 lx in the winter, and significant seasonal differences under red light (P <0.05) and blue light (P <0.01) were also observed at 1000 lx.
The present study demonstrated that dynamic changes in the physiological effects of colors of light on autonomic functions via the non-visual pathway may be associated with the temperature regulation system.
PMCID: PMC3686708  PMID: 23587209
Body temperature regulation; Shivering; Sweating; Core interthreshold zone; Color of light

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