Photosynthetic acclimation of plants to growth irradiance: the relative importance of specific leaf area and nitrogen partitioning in maximizing carbon gain

Changes in specific leaf area (SLA, projected leaf area per unit leaf dry mass) and nitrogen partitioning between proteins within leaves occur during the acclimation of plants to their growth irradiance. In this paper, the relative importance of both of these changes in maximizing carbon gain is quantified. Photosynthesis, SLA and nitrogen partitioning within leaves was determined from 10 dicotyledonous C₃ species grown in photon irradiances of 200 and 1000 µmol m^?2 s^?1. Photosynthetic rate per unit leaf area measured under the growth irradiance was, on average, three times higher for high‐light‐grown plants than for those grown under low light, and two times higher when measured near light saturation. However, light‐saturated photosynthetic rate per unit leaf dry mass was unaltered by growth irradiance because low‐light plants had double the SLA. Nitrogen concentrations per unit leaf mass were constant between the two light treatments, but plants grown in low light partitioned a larger fraction of leaf nitrogen into light harvesting. Leaf absorptance was curvilinearly related to chlorophyll content and independent of SLA. Daily photosynthesis per unit leaf dry mass under low‐light conditions was much more responsive to changes in SLA than to nitrogen partitioning. Under high light, sensitivity to nitrogen partitioning increased, but changes in SLA were still more important.     

Leaf Structure of Tobacco In Vitro Grown Plantlets as Affected by Saccharose and Irradiance

Tobacco plantlets were cultured in vitro under high (200 µmol m^–2 s^–1) or low (60 µmol m^–2 s^–1) irradiance with or without saccharose in the medium. Light microscopy and image analysis were used to evaluate the effect of these culture conditions on leaf anatomy. Addition of saccharose resulted in thicker leaves (all leaf layers) and larger mesophyll cells under both growth irradiances. Various irradiance affected leaf anatomy differently when plantlets had been cultivated in presence or absence of saccharose in the medium. While under high irradiance in presence of saccharose leaf thickness and number of chloroplasts per cell section were increased, plantlets grown under high irradiance in absence of saccharose had thinner leaves and less chloroplasts per cell section. The changes were more pronounced in palisade parenchyma layer.     

Response of plasmodesmata formation in leaves of C4 grasses to growth irradiance

Rapid metabolite diffusion across the mesophyll (M) and bundle sheath (BS) cell interface in C₄ leaves is a key requirement for C₄ photosynthesis and occurs via plasmodesmata (PD). Here, we investigated how growth irradiance affects PD density between M and BS cells and between M cells in two C₄ species using our PD quantification method, which combines three‐dimensional laser confocal fluorescence microscopy and scanning electron microscopy. The response of leaf anatomy and physiology of NADP‐ME species, Setaria viridis and Zea mays to growth under different irradiances, low light (100 μmol m^?2 s^?1), and high light (1,000 μmol m^?2 s^?1), was observed both at seedling and established growth stages. We found that the effect of growth irradiance on C₄ leaf PD density depended on plant age and species. The high light treatment resulted in two to four‐fold greater PD density per unit leaf area than at low light, due to greater area of PD clusters and greater PD size in high light plants. These results along with our finding that the effect of light on M‐BS PD density was not tightly linked to photosynthetic capacity suggest a complex mechanism underlying the dynamic response of C₄ leaf PD formation to growth irradiance.     

Effects of irradiance on growth,photosynthesis, and water use efficiency of seedlings of the chaparral shrub,Ceanothus megacarpus

Summary The effects of irradiance during growth on biomass allocation, growth rates, leaf chlorophyll and protein contents, and on gas exchange responses to irradiance and CO₂ partial pressures of the evergreen, sclerophyllous, chaparral shrub, Ceanothus megacarpus were determined. Plants were grown at 4 irradiances for the growth experiments, 8, 17, 25, 41 nE cm^-2 sec^-1, and at 2 irradiances, 9 and 50 nE cm^-2 sec^-1, for the other comparisons.At higher irradiances root/shoot ratios were somewhat greater and specific leaf weights were much greater, while leaf area ratios were much lower and leaf weight ratios were slightly lower than at lower irradiances. Relative growth rates increased with increasing irradiance up to 25 nE cm^-2 sec^-1 and then leveled off, while unit leaf area rates increased steeply and unit leaf weight rates increased more gradually up to the highest growth irradiance.Leaves grown at 9 nE cm^-2 sec^-1 had less total chlorophyll per unit leaf area and more per unit leaf weight than those grown at 50 nE cm^-2 sec^-1. In a reverse of what is commonly found, low irradiance grown leaves had significantly higher chlorophyll a/b than high irradiance grown leaves. High irradiance grown leaves had much more total soluble protein per unit leaf area and per unit dry weight, and they had much higher soluble protein/chlorophyll than low irradiance grown leaves.High irradiance grown leaves had higher rates of respiration in very dim light, required higher irradiances for photosynthetic saturation and had higher irradiance saturated rates of photosynthesis than low irradiance grown leaves. CO₂ compensation irradiances for leaves of both treatments were very low, <5 nE cm^-2 sec^-1. Leaves grown under low and those grown under high irradiances reached 95% of their saturated photosynthetic rates at 65 and 85 nE cm^-2 sec^-1, respectively. Irradiance saturated rates of photosynthesis were high compared to other chaparral shrubs, 1.3 for low and 1.9 nmol CO₂ cm^-2 sec^-1 for high irradiance grown leaves. A very unusual finding was that leaf conductances to H₂O were significantly lower in the high irradiance grown leaves than in the low irradiance grown leaves. This, plus the differences in photosynthetic rates, resulted in higher water use efficiencies by the high irradiance grown leaves. High irradiance grown leaves had higher rates of photosynthesis at any particular intercellular CO₂ partial pressure and also responded more steeply to increasing CO₂ partial pressure than did low irradiance grown leaves. Leaves from both treatments showed reduced photosynthetic capability after being subjected to low CO₂ partial pressures (100 bars) under high irradiances. This treatment was more detrimental to leaves grown under low irradiances.The ecological implications of these findings are discussed in terms of chaparral shrub community structure. We suggest that light availability may be an important determinant of chaparral community structure through its effects on water use efficiencies rather than on net carbon gain.     

INORGANIC CARBON REPLETION CONSTRAINS STEADY‐STATE LIGHT ACCLIMATION IN THE CYANOBACTERIUM SYNECHOCOCCUS ELONGATUS1

Cyanobacteria show high metabolic plasticity by re‐allocating macromolecular resources in response to variations in both environmental inorganic carbon (Ci) and light. We grew cultures of the picoplanktonic cyanobacterium Synechococcus elongatus Nägeli across a 50‐fold range of growth irradiance at either a dissolved [Ci] <0.1 mM, sufficient to induce strongly the carbon‐concentrating mechanism (CCM) or a dissolved [Ci] of ～4 mM, sufficient to strongly induce the CCM to basal constitutive activity. There was no detectable growth cost of acclimation to low Ci across the entire range of irradiance and growth was nearly light saturated at 50 l mol photons·m^?2·s^?1. Cells acclimated to low Ci significantly re‐allocated macromolecular resources to support their CCM, while maintaining near homeostatis of metabolic flux per unit photosynthetic complex. Changing growth irradiance also drove re‐organization of the photosynthetic machinery to balance excitation flux and metabolic demands, but flux per complex varied widely across the range of tolerable growth irradiances. Across the range of growth irradiance, low Ci cells had significantly less phycocyanin than high Ci cells, which corresponded to a lower PSII absorbance capacity. Furthermore, low Ci cells maintained more PSI per cell^?1 than high Ci cells under high growth irradiance. Low Ci cells could therefore maintain more of their PSII reaction centers open at high growth irradiance than could high Ci cells, which experienced a significant PSII closure. Thus, acclimation to growth under high available Ci actually constrained acclimation to high light by restricting electron transport downstream from PSII in S. elongatus.     

Shade adaptation of photosynthesis in Coffea arabica

The effect of irradiance on the rate of net photosynthesis was measured for mature leaves of coffee grown under five levels of radiation from 100% to 5% daylight. The rate of light-saturated photosynthesis per unit leaf area (P_Nmax) increased from 2 mol CO₂ m^-2 s^-1 under 5% daylight to 4.4 mol CO₂ m^-2 s^-1 under 100% daylight. The photon flux density (PAR, photosynthetically active radiation) needed for 50% saturation of photosynthesis, as well as the light compensation point, also increased with increasing levels of irradiation during growth. The quantum efficiency of photosynthesis (), measured by the initial slope of the photosynthetic response to increasing irradiance, was greater under shaded growth conditions. The rate of dark respiration was greatest for plants grown in full daylight. On the basis of the increase in the quantal efficiency of photosynthesis and the low light compensation point when grown under shaded conditions, coffee shows high shade adaptation. Plants adjusted to shade by an increased ability to utilize short-term increases in irradiance above the level of the growth irradiance (measured by the difference between photosynthesis at the growth irradiance, P_Ng, and P_Nmax).     

Dense crowns of the hemiparasitic mistletoe Viscum album L. exhibit shrub-like growth and high dry matter turnover

Growth characteristics and crown parameters of the evergreen, hemiparasitic mistletoe Viscum album were studied using a growth model based on the regular dichotomous growth and conservancy of branching patterns. Data were collected from one final harvest of 13 individuals of three different subspecies growing on the hosts Betula pendula, Abies alba and Pinus sylvestris. Compared to Viscum album ssp. album growing on deciduous hosts, the two subspecies abietis and austriacum (syn. laxum) on evergreen hosts showed longer average leaf life spans, but leaves continuously increased in area over time up to an age of at least 4 years irrespective of host. Relative growth rates in plant dry matter (RGR; 0.53–0.61 g g⁻¹ a⁻¹) were in the same range as of deciduous shrubs in central Europe but clearly higher than in mature forest trees. However, this growth is hidden within the spherical crowns of mistletoes, as annual increase in crown volume is extraordinarily low, resulting in very high leaf area densities (LAD; 7.7–17.6 m² m⁻³), not found in any other woody species. Besides differences in leaf size and phenology, the three subspecies exhibit similar patterns of partitioning. As in other mistletoe species, significant parts of dry matter are regularly allocated into fruit production (10–15%). Regarding the generally lower net photosynthesis rates in mistletoes as compared to their hosts, it is questioned, whether photoautotrophic carbon gain of the leaves is sufficient to maintain the observed high relative growth rates.     

Far-red light inhibits growth but promotes carotenoid accumulation in the green microalga Dunaliella bardawil

It has been demonstrated that far-red light reduces growth of marine phytoplankton and that light quality controls growth and photosynthetic metabolism in algae. The green halotolerant microalga, Dunaliella bardawil, accumulates high amounts of β-carotene (up to 10% of its dry weight) under conditions of high light or nutrient limitation. The influence of increasing irradiance and of far-red light in D. bardawil was studied. Continuous irradiance was provided by white fluorescent lamps alone (WL) or supplemented with far-red Linestra lamps (WL+FR). For both types of light, cultures were acclimatized at increasing irradiances (50-300 µmol m^?2 s^?1), and cell density, photosynthetic activity and pigment content were determined. Cell density increased with the photon irradiance, and was higher in WL than in WL+FR under the same irradiance, but the reverse occurred in respect of cell volume. Growth rate was higher under WL+FR. Far-red light induced faster growth but reduced the maximal cell density of the cultures. Chlorophyll a concentration was higher in white light, but total carotenoid content increased dramatically in both far-red light treatments (about 50% on a per cell basis) and with the increase of irradiance. Our results show that far-red light has a significant influence on growth and photosynthesis of D. bardawil, inducing a decrease in cell density, photosynthetic activity and chlorophyll concentration, and an increase in growth rate, cell volume and carotenoid content.     

Effects of Irradiance during Growth on Adaptive Photosynthetic Characteristics of Velvetleaf and Cotton

We grew velvetleaf (Abutilon theophrasti Medic.) and cotton (Gossypium hirsutum L. var. Stoneville 213) at three irradiances and determined the photosynthetic responses of single leaves to a range of six irradiances from 90 to 2000 μeinsteins m⁻²sec⁻¹. In air containing 21% O₂, velvetleaf and cotton grown at 750 μeinsteins m⁻²sec⁻¹ had maximum photosynthetic rates of 18.4 and 21.9 mg of CO₂ dm⁻²hr⁻¹, respectively. Maximum rates for leaves grown at 320 and 90 μeinsteins m⁻²sec⁻¹ were 15.3 and 10.3 mg of CO₂ dm⁻²hr⁻¹ in velvetleaf and 12 and 6.7 mg of CO₂ dm⁻²hr⁻¹ in cotton, respectively. In 1 O₂, maximum photosynthetic rates were 1.5 to 2.3 times the rates in air containing 21% O₂, and plants grown at medium and high irradiance did not differ in rate. In both species, stomatal conductance was not significantly affected by growth irradiance. The differences in maximum photosynthetic rates were associated with differences in mesophyll conductance. Mesophyll conductance increased with growth irradiance and correlated positively with mesophyll thickness or volume per unit leaf area, chlorophyll content per unit area, and photosynthetic unit density per unit area. Thus, quantitative changes in the photosynthetic apparatus help account for photosynthetic adaptation to irradiance in both species. Net assimilation rates calculated for whole plants by mathematical growth analysis were closely correlated with single-leaf photosynthetic rates.     

Structural and physiological sexual dimorphism estimated from three‐dimensional virtual trees of yerba‐mate (Ilex paraguariensis) is modified by cultivation environment

Yerba‐mate is a subtropical, evergreen, dioecious, South American tree. Sexual dimorphism in photosynthesis, leaf allometry and foliage distribution was hypothesised. Virtual trees (constructed in VPlants software from detailed measurements of plant morphogenesis) of the two genders were compared considering two contrasted cultivation environments and three developmental stages. The total crown volume, leaf area per plant (LA), leaf area index (LAI) and leaf area density (LAD) were calculated. The light interception and photosynthesis were computed from mock‐ups in VegeSTAR. Structural sexual dimorphism concerned general plant form, internode length, leaf allometry, leaf surface, pattern of leaf area distribution and LAD. Cultivation environment and developmental stage acted strongly on sex expression of all observed structural parameters and physiological stages. Sexual differentiation in LA and light interception was related to leaves positioned in the lowest layers (150 cm above ground), whereas sexual specialisation in leaf and plant photosynthesis was related to early vegetative and reproductive stages. Several sexual responses strongly depended on the environment, especially light conditions, with opposite effects observed on female and male plants whether they were cultivated in monoculture or in forest understorey, under high‐light condition or low‐light condition, respectively. Optimised foliage structure and physiology in females may compensate for greater reproductive costs in early developmental stages, but females and males equalise in photosynthetic efficiency after 2‐year regrowth.     

Effect of irradiance during growth of Glycine max on photosynthetic capacity and percent activation of ribulose 1,5-bisphosphate carboxylase

The initial (in vivo) and total (activity present after preincubation with CO₂ and Mg²⁺) activities of ribulose bisphosphate carboxylase were both assayed in extracts of leaves of soybean (Glycine max) plants which had been grown under 4 different irradiance levels. The total carboxylase activity per unit leaf area decreased with decreased irradiance during growth but was not different on a dry weight basis. The initial activity as a percentage of the total activity was unchanged (approximately 95%) except in leaves of plants grown at the lowest irradiance (74%). When the plants grown at the lowest irradiance were exposed to high irradiance, the initial activity was increased to 93% of the total. Light saturated rates of photosynthesis per unit leaf area were lower and saturated at lower irradiance for plants grown at lower irradiances. Initial carboxylase activity was correlated closely (r²=0.84) with leaf photosynthesis rate on a dry weight basis.     

Carbon gain and photosynthetic response of chrysanthemum to photosynthetic photon flux density cycles

Most models of carbon gain as a function of photosynthetic irradiance assume an instantaneous response to increases and decreases in irradiance. High- and low-light-grown plants differ, however, in the time required to adjust to increases and decreases in irradiance. In this study the response to a series of increases and decreases in irradiance was observed in Chrysanthemum × morifolium Ramat. “Fiesta” and compared with calculated values assuming an instantaneous response. There were significant differences between high- and low-light-grown plants in their photosynthetic response to four sequential photosynthetic photon flux density (PPFD) cycles consisting of 5-minute exposures to 200 and 400 micromoles per square meter per second (μmol m⁻²s⁻¹). The CO₂ assimilation rate of high-light-grown plants at the cycle peak increased throughout the PPFD sequence, but the rate of increase was similar to the increase in CO₂ assimilation rate observed under continuous high-light conditions. Low-light leaves showed more variability in their response to light cycles with no significant increase in CO₂ assimilation rate at the cycle peak during sequential cycles. Carbon gain and deviations from actual values (percentage carbon gain over- or underestimation) based on assumptions of instantaneous response were compared under continuous and cyclic light conditions. The percentage carbon gain overestimation depended on the PPFD step size and growth light level of the leaf. When leaves were exposed to a large PPFD increase, the carbon gain was overestimated by 16 to 26%. The photosynthetic response to 100 μmol m⁻² s⁻¹ PPFD increases and decreases was rapid, and the small overestimation of the predicted carbon gain, observed during photosynthetic induction, was almost entirely negated by the carbon gain underestimation observed after a decrease. If the PPFD cycle was 200 or 400 μmol m⁻² s⁻¹, high- and low-light leaves showed a carbon gain overestimation of 25% that was not negated by the underestimation observed after a light decrease. When leaves were exposed to sequential PPFD cycles (200-400 μmol m⁻² s⁻¹), carbon gain did not differ from leaves exposed to a single PPFD cycle of identical irradiance integral that had the same step size (200-400-200 μmol m⁻² s⁻¹) or mean irradiance (200-300-200 μmol m⁻² s⁻¹).     

不同密度红桦幼苗苗冠结构与竞争对CO2浓度升高的响应

研究了两个种植密度下,红桦 (Betula albosinensis)苗冠结构特征对CO₂浓度的响应,在此基础上探讨了CO₂浓度升高对植物竞争压力的影响。结果表明,冠幅、冠高、苗冠表面积和苗冠体积均受CO₂浓度升高的影响而增加,但是受密度增加的影响而降低。CO₂浓度升高对苗冠的促进效应在低密度条件下大于高密度处理,高密度条件下苗冠基本特征部分地受到CO₂浓度升高的促进作用;升高种植密度的效应则在高CO₂浓度条件下大于现行CO₂浓度处理。高CO₂浓度和高密度条件下,LDcpa（单位苗冠投影面积叶片数）、LDcv（单位苗冠体积叶片数） 和苗冠底部枝条的枝角均低于相应的现行CO₂浓度处理和低密度处理,这主要是由于冠幅和冠高的快速生长所造成的。升高CO₂浓度对枝条长度的影响与枝条在主茎上所处位置有关。总之,升高CO₂浓度有利于降低增加种植密度对苗冠所带来的负效应,而增加种植密度降低了升高CO₂浓度的正效应。LDcpa和LDcv的降低表明,红桦在升高CO₂浓度和种植密度的条件下,会作出积极的响应,从而缓解由于生长的增加所带来的竞争压力的增加。     

Effects of daylength,irradiance and settlement density on the growth and reproduction of <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> gametophytes

The effects of daylength, irradiance and spore settlement density on the growth, maturation and sporophyte production of Undaria pinnatifida (Harvey) Suringar gametophytes were examined using a factorial experimental design in culture. The growth of Undaria gametophytes increased with increasing daylength (8, 12 and 16 h), but the maximum fertility occurred at a daylength of 12 h followed by 8 and 16 h. Gametophytes grew better at the 16 h daylength under the same mean daily irradiance (MDI) of 20 μmol photons m⁻² s⁻¹. However, the fertility was higher at the short daylength (8 h), indicating that the maturation of U. pinnatifida gametophytes is influenced by daylength rather than by the MDI. Vegetative growth and sporophyte production of gametophytes were better at 60 μmol photons m⁻² s⁻¹ than at 30 μmol photons m⁻² s⁻¹ under a 8:16 h LD (Light: Dark) cycle, and their growth and maturation were density-dependant in 16 and 12 h daylength, respectively. These results suggest that the U. pinnatifida gametophytes require a certain amount of light for the growth and reproduction, and intraspecific competition occurred under the optimal growth and maturation conditions. However, the sporophyte recruits per unit has been enhanced with increasing spore settlement density at 8 and 12 h daylengths indicating that high settlement density gives a benefit for maintaining population, even though the sporophyte production of each female plant is inhibited. In conclusion, the vegetative growth, reproduction and sporophyte production of U. pinnatifida gametophytes are retarded at a low irradiance above growth saturation and a high settlement density, and are determined by daylength.     

不同光环境下紫椴幼树树冠结构的可塑性响应

从冠形、侧枝和叶片在树冠中的空间分布角度对天然更新紫椴幼树的树冠结构进行了论述,认为紫椴幼树树冠对光照条件的变化有显著的可塑性响应.强光通过抑制主干的生长促进了侧枝的分化,庇荫则通过抑制1级侧枝的生长促进了侧枝的再分枝.随着光照水平的降低,紫椴幼树的数量叶片密度显著降低,且叶片逐渐集中于冠上层.林冠下的紫椴幼树通过这种侧枝和叶片的分布格局,在形态上提高其对光的截获能力在适度庇荫环境中,紫椴幼树垂直生长采取演替先锋种的"避荫”对策,侧枝生长采取中等耐荫种的"掠光”对策;在弱光环境中,紫椴幼树则采取典型的忍耐适应行为.这种树冠结构的变化是提高紫椴幼树对光的截获能力的一种有益适应.     

Activities of Noncyclic and Alternative Pathways of Photosynthetic Electron Transport in Leaves of Broad Bean Plants Grown at Various Light Irradiances

Activities of noncyclic and alternative pathways of photosynthetic electron transport were studied in intact leaves of broad been (Vicia faba L.) seedlings grown under white light at irradiances of 176, 36, and 18 µmol quanta/(m² s). Electron flows were followed from light-induced absorbance changes at 830 nm related to redox transformations of P700, the photoactive PSI pigment. The largest absorbance changes at 830 nm, induced by either white or far-red light, were observed in leaves of seedlings grown at irradiance of 176 µmol quanta/(m² s), which provides evidence for the highest concentration of PSI reaction centers per unit leaf area in these seedlings. When actinic white light of 1800 µmol quanta/(m² s) was turned on, the P700 oxidation proceeded most rapidly in leaves of seedlings grown at irradiance of 176 µmol quanta/(m² s). The rates of electron transfer from PSII to PSI were measured from the kinetics of dark P700⁺ reduction after turning off white light. These rates were similar in leaves of all light treatments studied, and their characteristic reaction times were found to range from 9.2 to 9.5 ms. Four exponentially decaying components were resolved in the kinetics of dark P700⁺ reduction after leaf exposure to far-red light. A minor but the fastest component of P700⁺ reduction with a halftime of 30–60 ms was determined by electron transfer from PSII, while the three other slow components were related to the operation of alternative electron transport pathways. Their halftimes and relative magnitudes were almost independent on irradiance during plant cultivation. It is concluded that irradiance during plant growth affects the absolute content of PSI reaction centers in leaves but did not influence the rates of noncyclic and alternative electron transport.From Fiziologiya Rastenii, Vol. 52, No. 4, 2005, pp. 485–491.Original English Text Copyright © 2005 by Nikolaeva, Bukhov, Egorova.The article was translated by the authors.     

The responses of light interception,photosynthesis and fruit yield of cucumber to LED‐lighting within the canopy

Mathematical models of light attenuation and canopy photosynthesis suggest that crop photosynthesis increases by more uniform vertical irradiance within crops. This would result when a larger proportion of total irradiance is applied within canopies (interlighting) instead of from above (top lighting). These irradiance profiles can be generated by Light Emitting Diodes (LEDs). We investigated the effects of interlighting with LEDs on light interception, on vertical gradients of leaf photosynthetic characteristics and on crop production and development of a greenhouse‐grown Cucumis sativus‘Samona’ crop and analysed the interaction between them. Plants were grown in a greenhouse under low natural irradiance (winter) with supplemental irradiance of 221 µmol photosynthetic photon flux m^?2 s^?1 (20 h per day). In the interlighting treatment, LEDs (80% Red, 20% Blue) supplied 38% of the supplemental irradiance within the canopy with 62% as top lighting by High‐Pressure Sodium (HPS)‐lamps. The control was 100% top lighting (HPS lamps). We measured horizontal and vertical light extinction as well as leaf photosynthetic characteristics at different leaf layers, and determined total plant production. Leaf mass per area and dry mass allocation to leaves were significantly greater but leaf appearance rate and plant length were smaller in the interlighting treatment. Although leaf photosynthetic characteristics were significantly increased in the lower leaf layers, interlighting did not increase total biomass or fruit production, partly because of a significantly reduced vertical and horizontal light interception caused by extreme leaf curling, likely because of the LED‐light spectrum used, and partly because of the relatively low irradiances from above.     

Light-Induced Adaptive Responses under Greenhouse and Controlled Conditions in the Fern Pteris cretica var. ouvrardii

The influence of different growth irradiance conditions on plant development and foliar features were assessed in a shade-adapted fern variety, Pteris cretica var. ouvrardii. A comparison of frond morphology, anatomy, chloroplast infrastructure and chlorophyll content of plants cultivated in both greenhouse and in controlled growth chambers under moderate light, low light (control) and extreme shade revealed pronounced phenotypic modifications. Moderate light induced decreased frond surfaces, thicker leaves, lower chlorophyll content per surface unit, and a markedly reduced density of intraplastidial membranes. In contrast, morphological responses to extreme shade included the formation of larger, thinner fronds, increased chlorophyll content; and a higher membrane density in chrloplasts. A dorsi-ventral distribution of starch-gorged chlroplasts (lower mesophyll cell layers) and essentially starch-free chloroplasts (upper cell layer) characterizes low-light and moderate light fronds, while homogenous starch-free chloroplasts are present in all cell layers of extreme shade fronds. The light-induced modifications are discussed as adaptive responses.     

Effect of irradiance on shoot development in vitro

The growth of Disanthus cercidifolius and Rhododendron cultivars, and to a lesser extent Crataegus oxyacantha cv. Paul's Scarlet, was modified by altering either the spectral quality or the level of irradiance received by shoot cultures; which were otherwise maintained under uniform medium and plant growth regulator (PGR) conditions in vitro. When the spectrum of Philips colour 84 (white) fluorescent lamps was modified by coloured cellulose acetate filters, red light promoted shoot extension and axillary branching, whereas blue light inhibited shoot growth and reduced leaf chlorophyll content in the sensitive cultivar R. cv. Dopey. By using single or multiple layers of neutral density filters, or moving cultures nearer to the light source, irradiance from white light was varied. All cultures grew well at low levels of irradiance (c. 11 µmol m^–2 s^–1), but the growth and leaf chlorophyll content of cultures of Disanthus and Rhododendron cultivars were suppressed by increasing irradiance. In three related Rhododendron cultivars, increased irradiance promoted the development of adventitious shoots. Crataegus shoots were tolerant of a wide range of irradiances and only shoot extension was inhibited at the highest level tested; leaf chlorophyll content was unaffected. These results are discussed in terms of the differential perception of light relative to the natural habitats of these plants, and of the possible direct effect of irradiance upon PGRs in the culture system.