A dynamic analysis of the shade-induced plasticity in Arabidopsis thaliana rosette leaf development reveals new components of the shade-adaptative response

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.     

Correlation between leaf growth variables suggest intrinsic and early controls of leaf size in Arabidopsis thaliana

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.     

Plant development controls leaf area expansion in alfalfa plants competing for light

Background and Aims

The growth of crops in a mixture is more variable and difficult to predict than that in pure stands. Light partitioning and crop leaf area expansion play prominent roles in explaining this variability. However, in many crops commonly grown in mixtures, including the forage species alfalfa, the sensitivity and relative importance of the physiological responses involved in the light modulation of leaf area expansion are still to be established. This study was designed to assess the relative sensitivity of primary shoot development, branching and individual leaf expansion in alfalfa in response to light availability.

Methods

Two experiments were carried out. The first studied isolated plants to assess the potential development of different shoot types and growth periods. The second consisted of manipulating the intensity of competition for light using a range of canopies in pure and mixed stands at two densities so as to evaluate the relative effects on shoot development, leaf growth, and plant and shoot demography.

Key Results

Shoot development in the absence of light competition was deterministic (constant phyllochrons of 32·5 °Cd and 48·2 °Cd for primary axes and branches, branching probability of 1, constant delay of 1·75 phyllochron before axillary bud burst) and identical irrespective of shoot type and growth/regrowth periods. During light competition experiments, changes in plant development explained most of the plant leaf area variations, with average leaf size contributing to a lesser extent. Branch development and the number of shoots per plant were the leaf area components most affected by light availability. Primary axis development and plant demography were only affected in situations of severe light competition.

Conclusions

Plant leaf area components differed with regard to their sensitivity to light competition. The potential shoot development model presented in this study could serve as a framework to integrate light responses in alfalfa crop models.     

Leaf expansion and cell division are affected by reducing absorbed light before but not after the decline in cell division rate in the sunflower leaf

The effect of absorbed photosynthetic photon flux density (PPFD) on leaf expansion is a key issue for analysing the phenotypic variability between plants and for modelling feedback loops. Expansion and epidermal cell division in leaf 8 of sunflower were analysed in a series of five experiments where absorbed photosynthetic photon flux density (PPFD) was reduced either by shading or by covering part of the leaf area. These treatments were imposed at different times during leaf development. Expansion and cell division were affected by a reduction in absorbed PPFD only in the first part of leaf development, while the leaf area was less than 2% of its final value and while absolute expansion rate was slow. In contrast, it was not affected if imposed later when the leaf was visible and absolute expansion rate was at maximum. A reduction in absorbed PPFD caused the same reduction in expansion and in cell division whether it was due to a reduction in incident PPFD or to a reduction in photosynthetic leaf area, suggesting that carbon metabolism was involved. Relative expansion rate recovered to control levels when relative division rate began to decline, in all experiments and in all zones of a leaf. This was probably linked to the source–sink transition, after which the leaf had such a high priority in carbon allocation that it was largely insensitive to changes in absorbed PPFD. The final leaf area was therefore closely related to the cumulated PPFD absorbed by the plant from leaf initiation to the end of exponential cell division.     

<Emphasis Type="Italic">ATHB23</Emphasis>, an <Emphasis Type="Italic">Arabidopsis</Emphasis> class I homeodomain-leucine zipper gene,is expressed in the adaxial region of young leaves

Homeobox genes are essential regulators of plant development. ATHB23, a class I homeodomain leucine zipper gene of Arabidopsis, was found to be induced by treatment with the phytohormone gibberellin (GA). In order to clarify its role in development, we performed a histochemical analysis of transgenic plants containing a construct with a GUS::GFP reporter under the control of the 1.5 kb upstream region of ATHB23. The construct was mainly expressed in young leaves and the styles of flowers but not in mature leaves. Microscopic examination of young leaves revealed that it was expressed in the adaxial domain of leaf primordia and the rib meristem. Expression of ATHB23, like that of GA5 encoding GA 20-oxidase, was reduced in mutants related to adaxial-abaxial leaf polarity (phb-1d, se-2, and kan1 kan2). Reduced expression of the GUS::GFP reporter gene was also observed in an se-2 background. These results indicate that ATHB23 is under the control of GA and other activators such as PHB, and is involved in establishing polarity during leaf development.     

Chloroplast movement in Alocasia macrorrhiza

Chloroplast movements in a rainforest understory plant Alocasia macrorrhiza (L.) G. Don are striking, creating changes in leaf transmittance that are visible to the naked eye. We have characterized the light requirements for these changes and the resulting changes in light penetration to different cell layers within the leaf and through the entire thickness of the leaf. Plants were grown either in a relatively constant, growth-chamber environment or in a variable, greenhouse environment. Irradiance-response curves for chloroplast movement were the same for both groups of plants, saturating at about 1 000 μmol m⁻² s⁻¹, though only the greenhouse-grown plants normally encountered light sufficient to drive the movement. Chloroplast movement caused changes in whole-leaf transmittance on the order of a few percent across the entire visible spectrum. Transmittance changes were larger within the leaf, especially directly under the palisade layer. Chloroplast movement could be manipulated experimentally by removing blue wavelengths from the spectrum of incident light or by treating with cytochalasin D.     

Foliar growth of Eriocnema fulva Naudin (Melastomataceae) in a forest fragment in southeastern Brazil.

Eriocnema fulva Naudin is an endangered herbaceous, perennial, iteroparous, evergreen species geographically restricted to southeastern-center Minas Gerais State, SE Brazil. The individuals occur as patches on rocky riverbanks shaded by seasonal semideciduous Atlantic forest; they are fixed by roots and have a pending stem. Aiming to investigate leaf development and its importance for individual survival, fifteen contiguous plots (1 x 1 m) were set down in Jambreiro Forest (19 degrees 58'-59' S and 43 degrees 52'-55' W, 800-1100 m altitude), in the municipality of Nova Lima. A total of 260 individuals with the largest leaf blade length > or = 1 cm was tagged and measured in 1997, 1998, and 1999. Leaf expansion was recorded each month during 26 months until April 2000. Plant size was measured through leaf blade length, petiole length, stem length, and number of leaves. Significant changes were detected only after two years, thus indicating that plant growth is slow. The proportion of surviving leaves after two years was 60%. Total blade expansion took over 14 months, a slow growth rate when compared to leaves of other tropical forest canopy and understory species. Long leaf lifespans are to be found in plants exhibiting slow growth, and we observed that some leaves lived longer than three years. Petiole growth can help to better position the leaf in the search for light, thus contributing to the growth and survival of the plant. The relationships among size measures were significant, reinforcing the great contribution of leaf size for plant size. The age of the largest individual was estimated as 36 years based on the median annual leaf production rate.     

Leaf traits as indicators of limiting growing conditions for lettuce (Lactuca sativa)

Lettuce growth under unstressed conditions was compared to growth under four limiting conditions, i.e. no phosphorus fertilization (0_P), no nitrogen fertilization (0_N), low light (LR) and water stress (WR) over two different growing periods. We investigated the adaptive changes in terms of the morphological and physiological leaf traits, identifying stress‐specific and ‘stable’ indicators suitable for use in breeding programmes. The plants subjected to the WR treatments had lower leaf expansion and specific leaf area (SLA), as well as lower soil–plant analysis development (SPAD) values, stomatal conductance (POR), water index (WI) and leaf temperature (TIR) compared with plants in the unstressed CONTROL. Low light increased the leaf area (LA), SLA and leaf mass ratio (LMR). The 0_N treatment induced a general reduction in the normalised difference vegetation index (NDVI) values, as well as strong changes in LMR and SLA. In general, 0_P induced less pronounced effects than the other treatments. Principal component analysis indicated that the stable and suitable selection indicators of adaptive changes for low nitrogen and low light conditions were LA, SLA, leaf area per unit total plant mass (LAR), LMR, SPAD and POR, while SPAD, POR, TIR and WI were suitable indicators for drought.     

Two dominant photomorphogenic mutations of Arabidopsis thaliana identified as suppressor mutations of hy2

By screening suppressor mutants of the hy2 mutation of Arabidopsis thaliana , two dominant photomorphogenic mutants, shy1-1D and shy2-1D , for two genetic loci designated as SHY1 and SHY2 ( s uppressor of hy 2 mutation) have been isolated. Both of these non-allelic, extragenic suppressor mutations of hy2 are located on chromosome 1 of the Arabidopsis genome. Both mutations suppress the elongated hypocotyl phenotype of hy2 by light-independent inhibition of hypocotyl growth as well as by increasing the effectiveness of light inhibition of hypocotyl elongation. The shy1-1D mutation is partially photomorphogenic in darkness with apical hook opening and reduced hypocotyl elongation. The shy2-1D mutant displays highly photomorphogenic characteristics in darkness such as true leaf development, cotyledon expansion, and extremely reduced hypocotyl growth. In regard to hypocotyl elongation, however, the shy2-1D mutation is still light sensitive. Examination of red/far-red light responses shows that the shy1-1D mutation suppresses the hypocotyl elongation of the hy2 mutation effectively in red light but not effectively in far-red light. The shy2-1D suppresses hypocotyl elongation of the hy2 mutation effectively in both red and far-red light. Both mutations can also suppress the early-flowering phenotype of hy2 and have a distinct pleiotropic effect on leaf development such as upward leaf rolling. The data obtained suggest that SHY1 and SHY2 represent a novel class of components involved in the photomorphogenic pathways of Arabidopsis . This is the first report on the identification of dominant mutations in the light signal transduction pathway of plants.     

Plant leaf area expansion and assimilate production in wheat (Triticum aestivum L.) growing under low phosphorus conditions

Under phosphorus deficiency reductions in plant leaf area have been attributed to both direct effects of P on the individual leaf expansion rate and to a reduced availability of assimilates for leaf growth. In this work we use experimental and simulation techniques to identify and quantify these processes in wheat plants growing under P-deficient conditions. In a glasshouse experiment we studied the effects of soil P addition (0–138 kg P₂O₅ ha^-1) on tillering, leaf emergence, leaf expansion, plant growth, and leaf photosynthesis of wheat plants (cv. INTA Oasis) that were not water stressed. Plants were grown in pots containing a P-deficient (3 mg P g^-1 soil) sandy soil. Sowing and pots were arranged to simulate a crop stand of 173 plants m^-2. Experimental results were integrated in a simulation model to study the relative importance of each process in determining the plant leaf area during vegetative stages of wheat. Phosphorus deficiency significantly reduced plant leaf area and dry weight production. Under P-deficient conditions the phyllochron (PHY) was increased up to a 32%, compared to that of high-P plants. In low-P plants the rate of individual leaf area expansion during the quasi-linear phase of leaf expansion (LER) was significantly reduced. The effect of P deficiency on LER was the main determinant of the final size of the individual leaves. In recently expanded leaves phosphorus deficiency reduced the photosynthesis rate per unit leaf area at high radiation (AMAX), up to 57%. Relative values of AMAX showed an hyperbolic relationship with leaf P% saturating at 0.27%. Relative values of the tillering rate showed an hyperbolic relationship with the shoot P% saturating at values above 0.38%. The value of LER was not related to the concentration of P in leaves or shoots. A morphogenetic model of leaf area development and growth was developed to quantify the effect of assimilate supply at canopy level on total leaf area expansion, and to study the sensitivity of different model variables to changes in model parameters. Simulation results indicated that under mild P stress conditions up to 80% of the observed reduction in plant leaf area was due to the effects of P deficiency on leaf emergence and tillering. Under extreme P-deficient conditions the simulation model failed to explain the experimental results indicating that other factors not taken into account by the model, i.e. direct effects of P on leaf expansion, must have been active. Possible mechanisms of action of the direct effects of P on individual leaf expansion are discussed in this work.     

Changes in Leaf Phenology are Dependent on Tree Height inAcer mono,a Deciduous Broad-leaved Tree

Ontogenetic changes in leaf phenology of a hardwood tree,Acer mono, were investigated in individuals in different size classes in a temperate forest. Leaf emergence was earliest in current-year seedlings, and was increasingly delayed with increasing height of the individual. The shorter the tree, the longer the duration of leaf emergence. Timing of leaf emergence of the dominant heterospecific canopy trees was almost identical to that of conspecific adults; understorey light then gradually decreased with expansion of canopy leaves. These traits indicate that smaller individuals that receive the least light in summer can acquire favourable light for a longer period in spring than taller plants even in a forest understorey, but the advantage decreases with increasing plant height. Changes in the duration of leaf emergence and leaf longevity in response to environmental light regime [sun (forest edge)vs. shade (forest understorey)], were greatest for current year seedlings but decreased with increasing plant height. These results suggest that the plastic response of leaf phenology in juvenile stages may reduce the risk of losing an entire cohort in spatially heterogenous environments in the understorey of temperate forests.     

Leaf area expansion and assimilate production in sunflower (Helianthus annuus L.) growing under low phosphorus conditions

Reductions in leaf area and plant growth as a consequence of phosphorus (P) limitations have been attributed both to direct effects of P shortage on leaf expansion rate and to a reduced production of assimilates required for growth. Canopy assimilation and leaf area expansion are closely interrelated processes. In this work we used experimental and simulation techniques to identify and study their importance in determining leaf area on sunflower (Helianthus annuus L.) growing under P-deficient conditions. Experiment 1 was done outdoors, in Buenos Aires, Argentina, and Experiment 2 in a glasshouse in Wageningen, The Netherlands. In both experiments we studied the effects of soil P addition on leaf appearance, leaf expansion, dry matter accumulation, and leaf photosynthesis of non-water stressed plants grown in pots containing a P-deficient soil. Before sowing the equivalent amounts of 0–600 kg of super phosphate ha^-1 were added to the pots. Phosphorus deficiency delayed leaf appearance increasing the value of the phyllochron (PHY) up to 76%, the rate of leaf area expansion during the quasi-linear phase of leaf expansion (LER) was reduced by up to 74%, with respect to high P plants. Phosphorus deficiency reduced by up to 50% the rate of light saturated photosynthesis per unit of leaf area (AMAX) in recently expanded leaves, while at low levels of leaf insertion in the canopy, AMAX was reduced by up to 85%, when compared to that in high P plants. Phosphorus deficiency also reduced the duration of the quasi-linear phase of leaf expansion by up to eight days. The values of LER were related (r = 0.56, P < 0.05) to the mean concentration of P in all the leaves (Leaves P%) and not to the concentration of P in the individual leaf where LER was determined (r = 0.22, P < 0.4) suggesting that under P deficiency individual leaf expansion was not likely to be regulated by the total P concentration at leaf level. The values of AMAX of individual leaves were related (r = 0.79, P < 0.01) to the concentration of total P in the corresponding leaf (Leaf P%). LER showed a hyperbolic relationship with Leaves P% (R² = 0.94, P < 0.01, n = 13) that saturate at 0.14%. AMAX showed a hyperbolic relationship with Leaf P% (R² = 0.73, P < 0.01, n = 53) that saturated with values of Leaf P% higher than 0.22. A morphogenetic model of leaf area development and growth was developed to quantify the effect of assimilate supply at canopy level on total leaf area expansion, and to study the effects of model parameters on the growth of sunflower plants under P-deficient conditions. With this model we identified the existence of direct effects of P deficiency on individual leaf area expansion. However, we calculated that under mild P stress conditions up to 83% of the reduction in the observed leaf area was explained by the particular effects of P% on the rate of leaf appearance, on the duration of the linear period of leaf expansion, and on the value of AMAX. We also calculated that the effects of P deficiency on the value of AMAX alone, explained up to 41% of the observed reductions in total leaf area between the highest and the intermediate P level in Experiment 2. Possible mechanisms of action of the direct effects of P on individual leaf expansion are discussed in this paper.     

Individual leaf development in Arabidopsis thaliana: a stable thermal-time-based programme

In crop species, the impact of temperature on plant development is classically modelled using thermal time. We examined whether this method could be used in a non-crop species, Arabidopsis thaliana, to analyse the response to temperature of leaf initiation rate and of the development of two leaves of the rosette. The results confirmed the large plant-to-plant variability in the studied isogenic line of the Columbia ecotype: 100-fold differences in leaf area among plants sown on the same date were commonly observed at a given date. These differences disappeared in mature leaves, suggesting that they were due to a variability in plant developmental stage. The whole population could therefore be represented by any group of synchronous plants labelled at the two-leaf stage and followed during their development. Leaf initiation rate, duration of leaf expansion and maximal relative leaf expansion rate varied considerably among experiments performed at different temperatures (from 6 to 26 degrees C) but they were linearly related to temperature in the range 6-26 degrees C, with a common x-intercept of 3 degrees C. Expressing time in thermal time with a threshold temperature of 3 degrees C unified the time courses of leaf initiation and of individual leaf development for plants grown at different temperatures and experimental conditions. The two leaves studied (leaf 2 and leaf 6) had a two-phase development, with an exponential phase followed by a phase with decreasing relative elongation rate. Both phases had constant durations for a given leaf position if expressed in thermal time. Changes in temperature caused changes in both the rate of development and in the expansion rate which mutually compensated such that they had no consequence on leaf area at a given thermal time. The resulting model of leaf development was applied to ten experiments carried out in a glasshouse or in a growth chamber, with plants grown in soil or hydroponically. Because it predicts accurately the stage of development and the relative expansion rate of any leaf of the rosette, this model facilitates precise planning of sampling procedures and the comparison of treatments in growth analyses.     

Recovery of leaf area through accelerated shoot ontogeny in thrips-damaged cotton seedlings

BACKGROUND AND AIMS: Leaf area of cotton seedlings (Gossypium hirsutum) can be reduced by as much as 50 % by early season thrips infestations, but it is well documented that plants can regain the difference in leaf area once infestation ceases. The processes involved in the recovery have not been identified. Hypotheses include enhancement of the photosynthetic rate of the damaged leaves, more efficient leaf construction (i.e. more leaf area per unit of dry matter invested in new leaves), and more branching. METHODS: This 2-year field study examined these hypotheses and found that thrips-affected plants recovered from a 30 % reduction in total leaf area. During the recovery period, repeated measurements of gas exchange, leaf morphology and individual leaf areas at all nodes were made to assess their contribution to the recovery. KEY RESULTS: Recovery was not achieved through the previously proposed mechanisms. The pattern of nodal development indicated that the duration of leaf expansion of the smaller deformed leaves was shorter than that of control leaves, possibly because they had fewer cells. The production and expansion of healthy upper node leaves in thrips-affected plants could, therefore, begin sooner, about 1-2.5 nodes in advance of control plants. The proposed process of recovery was evident but weaker in the second year where thrips numbers were higher. CONCLUSIONS: It is concluded that thrips-affected plants overcame the leaf area disparity through an accelerated ontogeny of main stem leaves. By completing the expansion of smaller but normally functioning lower node leaves earlier, resources were made available to the unfolding of larger upper node leaves in advance of control plants. The generality of this mode of plant resistance in pest damage remains to be determined.     

Water relations and leaf expansion: importance of time scale

The role of leaf water relations in controlling cell expansion in leaves of water-stressed maize and barley depends on time scale. Sudden changes in leaf water status, induced by sudden changes in humidity, light and soil salinity, greatly affect leaf elongation rate, but often only transiently. With sufficiently large changes in salinity, leaf elongation rates are persistently reduced. When plants are kept fully turgid throughout such sudden environmental changes, by placing their roots in a pressure chamber and raising the pressure so that the leaf xylem sap is maintained at atmospheric pressure, both the transient and persistent changes in leaf elongation rate disappear. All these responses show that water relations are responsible for the sudden changes in leaf elongation rate resulting from sudden changes in water stress and putative root signals play no part. However, at a time scale of days, pressurization fails to maintain high rates of leaf elongation of plants in either saline or drying soil, indicating that root signals are overriding water relations effects. In both saline and drying soil, pressurization does raise the growth rate during the light period, but a subsequent decrease during the dark results in no net effect on leaf growth over a 24 h period. When transpirational demand is very high, however, growth-promoting effects of pressurization during the light period outweigh any reductions in the dark, resulting in a net increase in growth of pressurized plants over 24 h. Thus leaf water status can limit leaf expansion rates during periods of high transpiration despite the control exercised by hormonal effects on a 24 h basis.     

Effect of Verticillium dahliae on Photosynthesis, Leaf Expansion and Senescence of Field-grown Sunflower

On the basis of known sunflower (Helianthus annuus L.) responsesto soil water deficit, it is proposed that the effect of thefungus Verticillium dahliae Klebahn on plant leaf area precedesand is greater than its effect on leaf photosynthesis and stomatalconductance. To test this hypothesis, we measured shoot andleaf area growth, leaf photosynthetic rate, stomatal conductanceand disease symptoms in a field experiment including hybridsof high (Sankol) and low (Dekasol 3900) susceptibility to V.dahliae. Plants inoculated with V. dahliae and controls werecompared. We also investigated the effect of V. dahliae on keycomponents of plant leaf area, leaf expansion and senescence,in inoculated and control plants of Sankol and Toba, a hybridof intermediate susceptibility to V. dahliae. Reduction in plantleaf area caused by V. dahliae was first detected 31 d afterinoculation (DAI), when visual symptoms of disease in inoculatedplants were slight (Sankol) or absent (Dekasol 3900). Reductionin leaf photosynthesis was first observed 66 DAI; stomatal conductanceand leaf dark respiration were both unaffected by V. dahliaeduring the whole experiment. In comparison with controls, V.dahliae reduced seasonal duration of plant leaf area by 25%in Dekalb 3900 and by 55% in Sankol, whereas the average reductionin leaf photosynthetic rate was 9%. In correspondence with thereduction in leaf area duration, inoculation reduced shoot drymatter of mature Sankol by 50%. In both experiments, less leafexpansion accounted for most of the early reduction in plantleaf area; as the disease progressed, increasing senescencealso contributed to reduced plant leaf area. It is concludedthat the response of sunflower to V. dahliae resembled the responseof the plant to soil water deficit: (1) plant leaf area, ratherthan leaf photosynthetic rate, accounted for the reduction ingrowth in mass; and (2) reduced leaf expansion early in theseason and faster leaf senescence in older plants accountedfor the decrease in plant leaf area. Copyright 2000 Annals ofBotany Company Helianthus annuus, Verticillium dahliae, allometry, apical dominance, drought, leaf expansion, leaf senescence, photosynthesis, stomatal conductance, growth     

Improved growth and water use efficiency of cherry saplings under reduced light intensity

Cherry (Prunus avium L.) saplings were grown under natural sunlight (controls) or moderate shading (up to 30%, depending on the incident light intensity and the hour of the day). Reduced light intensity increased the dry mass of each of the plant components studied. Consequently, the total dry mass of shaded plants was significantly greater than that of controls at the end of the growing season. However, the diurnal trend in the level of photosynthesis (per unit of leaf area) of shaded plants was similar to the controls in August, but lower in September. As the growing season proceeded, reduced photosynthetic rates, thinner mesophyll and larger specific leaf area in the shaded plants indicated that leaf development had adapted to shaded conditions throughout the growing season. It is suggested that increased growth of shaded plants was caused by a higher initial relative growth rate and a greater whole-plant photosynthesis. Shading consistently reduced transpiration over the season, therefore improving water use efficiency of shaded leaves. Our results suggest that a moderate reduction in light intensity can be a useful method for improving growth and saving water in hot and dry environments.     

The effect of light quality on leaf production and development of in vitro-cultured plants of Alternanthera brasiliana Kuntze

We investigated the influence of light quality on the leaf development of Alternanthera brasiliana Kuntze (Amaranthaceae) grown in vitro. Growth parameters including specific leaf mass, thickness, and leaf density were lowest in plants grown under red light. Blue light induced the largest number of leaves/plant, and the largest thickness and area of the leafblade. Green and red lights induced the smallest leaf areas. The thickness of the abaxial-face epidermis and spongy parenchyma of the plants was significantly reduced in plants grown under red light. The thickness of the palisade parenchyma and upper epidermis were significantly increased in plants grown under blue light, compared to the other fluorescent-light treatments. The specific spectral band also influenced the differentiation of mesophyll cells. In the dark and under red light, the mesophyll was homogenous; and in the dark and under green light, the leaves were more compact. Under blue light, the cells displayed the characteristic palisade morphology. The results showed that the increase of a specific parenchyma type was related to a specific spectral band. All spectral-quality treatments reduced the numbers of stomata and trichomes. The results for green light were in some respects similar to those for red light, and in other respects similar to those for blue light, probably because phytochromes and cryptochromes are green-light receptors. This study indicated that Alternanthera plants have strong morphological plasticity induced by light. The results suggest that high-quality Alternanthera can be achieved by culturing the plants in vitro under a combination of blue and red light.     

Variations in Tiller Dynamics and Morphology in Lolium multiflorum Lam.Vegetative and Reproductive Plants as affected by Differences in Red/Far-Red Irradiation

Lolium multiflorum Lam, plants were grown in a growth room undertwo light sources with red/far-red ratios of 1·62 and0·84 but similar photosynthetically active radiation.In both situations the capacity to produce new tillers and thelight available per tiller decreased with canopy growth. Tilleringwas further reduced by the low red/far-red ratio while lightinterception and plant dry weight were unaffected by this treatment.In both reproductive and vegetative plants under the lower red/far-redratio the time between leaf expansion and the appearance ofa tiller in its axil was increased and the proportion of ‘maturebuds’ that developed was reduced. Irradiation with lowred/far-red advanced the reproductive development and increasedthe number of fertile tillers per plant. It also caused longerleaf sheaths, blades and reproductive shoots. The results suggestthat as canopy density increases the lower light interceptionper tiller and the photomorphogenic effect of low red/far-redratios may reduce the capacity to produce new tillers. Lolium multiflorum, Lam., annual ryegrass, tillering, tiller growth, leaf growth, flowering, light quality.