Effects of reduced irradiance on leaf morphology,photosynthetic capacity,and fruit yield in olive (<Emphasis Type="Italic">Olea europaea</Emphasis> L.)

One-year-old olive trees (cv. Koroneiki) were grown in plastic containers of 50 000 cm³ under full daylight and 30, 60, and 90 % shade for two years. The effects of shade on leaf morphology and anatomy, including stomatal density and chloroplast structure, net photosynthetic rate (P _N), stomatal conductance (g _s), and fruit yield were studied. Shade reduced leaf thickness due to the presence of only 1–2 palisade layers and reduced the length of palisade cells and spongy parenchyma. The number of thylakoids in grana as well as in stroma increased as shade increased, while the number of plastoglobuli decreased in proportion to the reduced photosynthetically active radiation (PAR). The higher the level of shade, the lower the stomatal and trichome density, leaf mass per area (ALM), g _s, and P _N. Shade of 30, 60, and 90 % reduced stomatal density by 7, 16, and 27 %, respectively, while the corresponding reduction in P _N was 21, 35, and 67 %. In contrast, chlorophyll a+b per fresh mass, and leaf width, length, and particularly area increased under the same shade levels (by 16, 33, and 81 % in leaf area). P _N reduction was due both to a decrease in PAR and to the morphological changes in leaves. The effect of shade was more severe on fruit yield per tree (32, 67, and 84 %) than on P _N indicating an effect on bud differentiation and fruit set. The olive tree adapts well to shade compared with other fruit trees by a small reduction in stomatal and trichome density, palisade parenchyma, and a significant increase in leaf area.     

Blue light inhibits stomatal development in soybean isolines containing kaempferol-3-O-2G-glycosyl-gentiobioside (K9), a unique flavonoid glycoside

Stomata have a fundamental role in controlling plant photosynthesis and transpiration, but very little is known about factors controlling stomatal differentiation and development. Lines of soybean that contain a specific flavonol glycoside, kaempferol‐3‐O‐2‐glycosyl‐gentiobioside (K9), as well as greatly reduced stomatal density, especially on the adaxial epidermis, have been identified. The specific effects of blue light photoreceptors on stomatal development in K9 lines and their isoline pairs containing no K9 were studied. Low irradiances of blue light (7% of total photosynthetically active radiation) added to high irradiances from low‐pressure sodium lamps strongly inhibited stomatal development on the adaxial epidermis of K9 lines, but not in isoline pairs differing putatively in only one gene and lacking K9. Overall, blue light slightly increased stomatal density on the abaxial epidermis in all isolines, demonstrating differential regulation of stomatal development in the upper and lower epidermis. Blue light also caused an increase in leaf area in all isolines, indicating that changes in stomatal density were not the non‐specific result of alterations in leaf area. Morphological studies revealed that the blue light‐induced reduction in stomatal density in K9 lines was due to reduced stomatal initiation as well as aborted or abnormal stomatal development. As the phytochrome photostationary state was kept constant, the results indicate that one or more blue light receptors are involved in the control of stomatal development. This system should be useful for the study of mechanisms controlling stomatal development, even if the photo‐inhibitory response is unique to K9 lines.     

Stomatal numbers of soybean and response to water stress

The relationship among stomatal density, photosynthetic rate, leaf conductance, plant growth, bean yield and kaempferol triglucoside (K9) in the leaves of soybean (Glycine max (L.) Merr.) was examined in two field tests. K9 in the leaves was associated with reduced stomatal density, reduced photosynthetic rate, reduced stomatal conductance, reduced plant weight and lower bean yield. Plants with high stomatal frequency (lacking K9) were better able to take advantage of increased water supply by increasing stomatal conductance (upper surface), transpiration and bean yield. Plants with low stomatal frequency (with K9) were unresponsive to irrigation and in this sense were more tolerant of water stress, but their overall yield was low.     

Stomatal and photosynthetic responses to shade in sorghum, soybean and eastern gamagrass

We studied photosynthetic and stomatal responses of grain sorghum ( Sorghum bicolor [L.] Moench cv. Pioneer 8500), soybean ( Glycine max L. cv. Flyer) and eastern gamagrass ( Tripsacum dactyloides L.) during experimental sun and shade periods simulating summer cloud cover. Leaf gas exchange measurements of field plants showed that short-term (5 min) shading of leaves to 300–400 μmol m⁻² s⁻¹ photosynthetic photon flux density reduced photosynthesis, leaf temperature, stomatal conductance, transpiration and water use efficiency and increased intercellular CO₂ partial pressure. In all species, photosynthetic recovery was delayed when leaves were reilluminated, apparently by stomatal closure. The strongest stomatal response was in soybean. Photosynthetic recovery was studied further with soybeans grown indoors (maximum photosynthetic photon flux density 1 200 μmol m⁻² s⁻¹). Plants grown indoors had responses to shade similar to those of field plants, except for brief nonstomatal limitation immediately after reillumination. These responses indicated the importance of the light environment during leaf development on assimilation responses to variable light, and suggested different limitations on carbon assimilation in different parts of the soybean canopy. Photosynthetic oxygen evolution recovered immediately upon reillumination, indicating that the light reactions did not limit soybean photosynthetic recovery. While shade periods caused stomatal closure and reduced carbon gain and water loss in all species, the consequences for carbon gain/water loss were greatest in soybean. The occurrence of stomatal closure in all three species may arise from their shared phenologies and herbaceous growth forms.     

Ultraviolet-B effects on stomatal density,water-use efficiency,and stable carbon isotope discrimination in four glasshouse-grown soybean (Glyicine max) cultivars

Interactions between UV-B radiation and drought stress have been studied but the underlying mechanisms have not been thoroughly investigated. We hypothesized that UV-B radiation would improve water-use efficiency (WUE) by its effects on epidermal development, specifically stomatal density, and leaf gas exchange. Four lines of soybean (Glycine max: Essex, Williams, OX921, and OX922) were grown for 28 days in a glasshouse with and without supplemental UV-B radiation levels of 13 kJ m⁻² biologically effective radiation. UV-B radiation increased phenolic content of leaves in all lines and reduced leaf area in all lines except Williams. Adaxial stomatal density was reduced in Essex, OX921, and OX922 but abaxial stomatal density was reduced only in OX922. Stomatal conductance was reduced in concert with stomatal density as was internal CO₂ concentration (C_i). Instantaneous WUE was increased in Essex, Williams, and OX922. Stable carbon isotope analysis showed similar trends in long-term WUE, but these were only statistically significant in OX922. These results suggest that some soybean cultivars may respond to increased levels of UV-B by increasing WUE and that this response could be manifested through changes in stomatal development and functioning. Field studies are needed to test this hypothesis and to further evaluate the role of the K9 gene in this response.     

Acclimation of Myrtus communis to contrasting Mediterranean light environments - effects on structure and chemical composition of foliage and plant water relations

Leaf anatomical and chemical characteristics, water relations and stomatal regulation were studied in the shrub Myrtus communis growing under two contrasting Mediterranean light environments (full light versus 30% of full light) during the spring-summer period. These studies aimed to assess plant response to the combined effects of light and water availability. Foliar morphology, anatomy and chemistry composition acclimated positively to light conditions. Leaves of sun-exposed plants were thicker (38.7%) than those of shaded plants, mainly due to increased palisade parenchyma thickness, had a higher nitrogen concentration and stomatal density than the shade ones, which maximized foliar area (>SLA) and Chl/N molar ratio to improve light interception. Chlorophyll concentration per leaf area (Chl(a)) was always higher in sun leaves while, as expressed on dry mass (Chl(m)), significant differences were only apparent in September, shade leaves presenting higher values. During the summer period Chl(a) and Chl(m) markedly declined in sun leaves and remained unchanged in shade ones. The ratio of chlorophyll a/b was not affected either by the light intensity or by the season. Shade leaves presented generally a higher concentration of soluble carbohydrates per dry mass. No significant differences in starch concentration were apparent between sun and shade leaves and a gradual depletion occurred during the water stress period. Maximum stomatal conductances correlated positively with predawn water potential. Throughout the season, sun plants always presented higher leaf conductance to water vapour and lower minimum leaf water potentials, indicating an interaction of light-environment on these water relation parameters. Stomatal closure constitutes a mechanism to cope with diurnal and seasonal water deficits, sun plants presenting a more efficient control of water losses during water deficiency period. In addition, both sun and shade plants evidenced leaf osmotic adjustment ability in response to water stress, which was greater in sun ones.     

Effects of Shading the First Leaf on Growth of Barley Plants: I. Long-term Experiments

Plants of barley were grown under controlled conditions andthe first or second leaves covered with tubular shades thusreducing the light intensity at the leaf surface to low levels.Expansion of the shaded leaves was not prevented, but appearanceof the next leaf but one and all subsequent leaves on the mainstemwas delayed by up to 3 days. Primordia of the first four leaveswere present in the dry grain. Shade treatment delayed slightlythe initiation of the eighth and subsequent leaves and transitionto the double ridge stage at the mainstem apex. Shading the first leaf caused a temporary reduction in the rateof dry-matter increase of plants, but after 14 days the ratewas similar to that of control plants. Smaller effects werefound when the second leaf was shaded. Dry-matter productionfollowed two logarithmic phases in the period prior to awn emergence,and rates for the whole plant and for plant parts were similarfor control and shaded plants. Thus, apart from the initialperturbation, shading had no effect on growth in terms of rateof dry-weight gain. Shade treatment did not affect weight per grain or numbers ofgrain per ear, but over-all yield of grain was significantlyreduced since shading delayed the appearance of tillers andalso reduced the number of tillers bearing grain. The effectof shade was especially marked on tillers originating on primarytillers. Similar qualitative effects on tiller development werefound in an experiment on wheat.     

Salinity tolerance of 'Valencia' orange trees on rootstocks with contrasting salt tolerance is not improved by moderate shade

The effects of shading in combination with salinity treatments were studied in citrus trees on two rootstocks with contrasting salt tolerance to determine if shading could reduce the negative effects of salinity stress. Well-nourished 2-year-old 'Valencia' orange trees grafted on Cleopatra mandarin (Cleo, relatively salt tolerant) or Carrizo citrange (Carr, relatively salt sensitive), were grown either under a 50% shade cloth or left unshaded in full sunlight. Half the trees received no salinity treatment and half were salinized with 50 mM Cl- during two 9 week salinity periods in the spring and autumn interrupted by an 11 week rainy period. The shade treatment reduced midday leaf temperature and leaf-to-air vapour pressure deficit regardless of salinity treatments. In non-salinized trees, shade increased midday CO2 assimilation rate (A(CO2)) and stomatal conductance, but had no effect on leaf transpiration (E(lf)). Shade also increased leaf chlorophyll and photosynthetic water use efficiency (A(CO2)/E(lf)) in leaves on both rootstocks and increased total plant dry weight in Cleo. The salinity treatment reduced leaf growth and leaf gas exchange parameters. Shade decreased Cl- concentrations in leaves of salinized Carr trees, but had no effect on leaf or root Cl- of trees on Cleo. There were no significant differences in leaf gas exchange parameters of shaded and unshaded salinized plants but the growth reduction from salinity stress was actually greater for shaded than for unshaded trees. Shaded trees on both rootstocks had higher leaf Na+ than unshaded trees after the first salinity period, and this shade-induced elevated leaf Na+ persisted after the second salinity period in trees on Carr. Thus, shading did not alleviate the negative effects of salinity on growth and Na+ accumulation.     

Leaf gas exchange of beech (Fagus sylvatica L.) seedlings in lightflecks: effects of fleck length and leaf temperature in leaves grown in deep and partial shade

Summary Responses of leaf gas exchange in shade and half-shade grown seedlings of the European beech, Fagus sylvatica L., to constant light conditions indicate different phases of photosynthetic induction: an immediate, a fast and a subsequent slow phase. The slow phase has both biochemical and stomatal components. The higher the induction, the higher the lightfleck utilization efficiency (LUE) attributable to a lightfleck. LUE can be higher than 100% compared to a theoretical instantaneous response. Lightfleck quantum yield (total carbon gain attributable to a lightfleck per incident quantum density in the fleck) is highest in short pulses of light. Post-illumination carbon gain initially increases with fleck length, levelling off above 20 s. The lower the induction, the longer carbon is fixed post-illuminatively (up to 84 s) but the less carbon is gained. Shade leaves are induced much faster than partial shade leaves. They utilize series of lightflecks to become fully induced, while half-shade (and sun) leaves depend on continuous high light. Half-shade leaves lose induction faster in low light between lightflecks. High as well as low temperatures strongly delay induction in half-shade but not in shade leaves. In general, shade leaves are much better adapted to the dynamic light environment of the forest understorey; however, their water-use efficiency during induction is lower.Dedicated to Prof. O. L. Lange on the occasion of his 65th birthday     

Stem growth habit affects leaf morphology and gas exchange traits in soybean

Backgrounds and Aims

The stem growth habit, determinate or indeterminate, of soybean, Glycine max, varieties affects various plant morphological and developmental traits. The objective of this study is to identify the effect of stem growth habit in soybean on the stomatal conductance of single leaves in relation to their leaf morphology in order to better understand the ecological and agronomic significance of this plant trait.

Methods

The stomatal conductance of leaves on the main stem was measured periodically under favourable field conditions to evaluate g_max, defined as the maximum stomatal conductance at full leaf expansion, for four varieties of soybean and their respective determinate or indeterminate near isogenic lines (NILs). Leaf morphological traits including stomatal density, guard cell length and vein density were also measured.

Key Results

The value of g_max ranged from 0·383 to 0·754 mol H₂O m⁻² s⁻¹ across all the genotypes for both years. For the four pairs of varieties, the indeterminate lines exhibited significantly greater g_max, stomatal density, numbers of epidermal cells per unit area and total vein length per unit area than their respective determinate NILs in both years. The guard cell length, leaf mass per area and single leaf size all tended to be greater in the determinate types. The variation of g_max across genotypes and years was well explained by the product of stomatal density and guard cell length (r = 0·86, P < 0·01).

Conclusions

The indeterminate stem growth habit resulted in a greater maximum stomatal conductance for soybean than the determinate habit, and this was attributed to the differences in leaf structure. This raises the further hypothesis that the difference in stem growth habit results in different water use characteristics of soybean plants in the field. Stomatal conductance under favourable conditions can be modified by leaf morphological traits.Key words: Soybean, Glycine max, stem growth habit, stomatal conductance, stomatal density, guard cell length, near isogenic lines     

土壤水分与短期遮光对棉花光合及其气孔响应的影响

以陆地棉（Ｇｏｓｓｙｐｉｕｍ ｈｉｒｓｕｔｕｍ Ｌ．Ｚｈｏｎｇｍａｉｎ,Ｎｏ２３）为供试材料,探讨了在充分供水－水分胁迫－复水的处理过程中,短期不同遮光水平对棉花光合特性及其气孔响应的影响。结果表明,在水分处理过程中,所有不同遮光水平的棉花叶片对短期遮光具有相似的基本响应规律;短期遮光使净光合速率迅速降低,气孔导度减少,但减少速率缓慢;遮阳网去掉后,叶片气 重新开放速率和光合恢复被延迟,水分胁迫期     

Do fruit sunburn control measures affect leaf photosynthetic rate and stomatal conductance in ‘Royal Gala’ apple?

A field study was conducted on a 5-year-old orchard of ‘Royal Gala’ apple (Malus domestica Borkh.) in Stellenbosch, South Africa, to investigate whether the measures employed to control sunburn in fruit, viz., evaporative cooling, Surround WP and 20% black shade net affect leaf photosynthetic gas exchange attributes in comparison to untreated control during the 2003/2004 season. Shade net significantly reduced midday leaf net photosynthetic rate (A) compared to evaporative cooling. Furthermore, shade net and Surround WP significantly reduced midday leaf stomatal conductance (g_s) compared to evaporative cooling and control. Evaporative cooling increased light saturated photosynthetic rate by 27 and 24% compared to shade net and Surround WP, respectively. Light compensation point and dark respiration of shaded leaves were about a third of the other treatments and about 50% less than the control leaves, respectively. Shade net down-regulated photosynthetic capacity of the leaves as evidenced by lower maximum rate of carboxylation and light saturated rate of electron transport compared to control leaves. Sunburn control treatments reduced day respiration by 60–70% compared to the control. Response of A and g_s to increasing temperature showed only slight increase in both A and g_s with increasing temperature from 20 to 30 °C. A declined at 35 °C in Surround WP and shade net leaves while it declined at 40 °C in evaporatively cooled and control leaves. Evaporative cooling and control had higher g_s than shade net and Surround WP at all leaf temperatures. In conclusion, shade net down-regulated photosynthetic reactions and Surround WP and shade net reduced leaf g_s and increased the vulnerability of leaf A and g_s to high temperature compared to evaporative cooling and control.     

七种热带雨林树苗叶片气孔特征及其可塑性对不同光照强度的响应

对生长在荫棚3种不同光照条件下和全自然光下的热带雨林4个冠层种(望天树、绒毛番龙眼、团花、红厚壳)和3个中层种(玉蕊、藤黄、滇南风吹楠)树苗叶片气孔特征以及它们的可塑性进行了研究、结果表明,这些植物的气孔全部着生在远轴面．7种植物中,玉蕊和绒毛番龙眼的气孔密度较大,滇南红厚壳和团花的保卫细胞最长．随光强的增大,气孔密度和气孔指数增大,单位叶气孔数在低光强下较大．除团花外,其它植物叶片气孔导度在50％光强处最大,而光强对保卫细胞的长度影响不显著．相关分析表明,气孔密度与植物单位叶的面积呈负相关。而与气孔导度的相关性不显著、尽管两种不同生活型植物气孔指数和单位叶气孔数在不同光强下的可塑性差异较小,但冠层树种气孔密度和气孔导度的可塑性显著高于中层树种．     

Photosynthesis in flashing light in soybean leaves grown in different conditions. I. Photosynthetic induction state and regulation of ribulose-1,5-bisphosphate carboxylase activity

The photosynthetic induction state under conditions of different lightfleck frequencies or durations, or different shade periods was studied in soybean leaves in order to examine how it might limit utilization of sunflecks in leaf canopies. Induction following an increase in photon flux density (PFD) from strongly limiting to saturating PFDs exhibited two phases; a fast-inducing one, requiring about 1 min and a slow one, requiring up to 60 min for completion. Transfer of fully induced leaves to low light resulted in a rapid decrease in the fast-inducing component, a slower decrease in the slow-inducing component and an even slower decrease in stomatal conductance. Therefore, the decreases in extent of induction appeared to be due to biochemical factors and not to stomatal closure. Under flashing light regimes consisting of 1-s lightflecks given at different frequencies for long periods, a constant induction state was achieved, the measure of induction state increased with the frequency of the lightflecks. This constant induction state also depended on the growth conditions, with shade leaves having a higher value than those grown at high light at any particular lightfleck frequency. The measure of induction state was mostly lower in flashing light as compared to constant light of the same mean PFD, particularly in leaves with a low light saturation point and in short lightflecks. Initial activities of ribulose-1,5-bisphosphate carboxylase (rubisco) were also higher in continuous light and were highly correlated with the measure of induction state. The rapid decrease in extent of induction of soybean leaves during shade periods is an important limitation to the ability of the leaves to respond to light increases similar to those occurring with sunflecks. At least part of the limitation on carbon assimilation during sunflecks due to photosynthetic induction is based on regulation of rubisco activity.     

Variations in stomatal density and index: implications for palaeoclimatic reconstructions

The variation in stomatal characters in leaves from one Alnus glutinosa (L.) Gaertn. tree is analysed. Measurements were taken from over 70 sites on the abaxial surfaces of representative ‘sun’ and ‘shade’ leaves having the same insertion point. The mean values of stomatal density and index in the shade leaf were significantly lower (71 and 93%, respectively) than those for the sun leaf. Within leaves, up to 2.5-fold differences in stomatal density values were observed. Contour maps derived from the data reveal non-random trends over the leaf surface. Correlations between stomatal density, epidermal cell density and stomatal index indicate that the variation in stomatal density within leaves arose primarily from local differences in stomatal differentiation, rather than from local differences in leaf expansion. This research demonstrates that a high level of variation in stomatal characters occurs both within and between leaves. We conclude that a well-defined sampling strategy should be used when estimating stomatal characters for (tree) leaves. Furthermore, the leaf's insertion point and situation within the tree crown should be taken into account. We discuss the implications of these findings for palaeoclimatic interpretations and emphasize the need for great caution when drawing conclusions based solely on stomatal characters.     

Effects of canopy shade on the lipid composition of soybean leaves

The effect of canopy shade on leaf lipid composition was examined in soybeans ( Glycine max cv. Young) grown under field conditions. Expanding leaves were tagged at 50, 58 and 65 days after planting (DAP) in plots with either a high (10 plants m⁻¹ row) or low (1 plant m⁻¹ row) plant density. At 92 DAP, light conditions ranged from a pho-tosynthetic photon flux density (PPFD) of 87% of full sun with a far-red/red (735 nm/645 nm) ratio of 0.9 at upper canopy leaves to extreme shade where the PPFD was 10% of full sun with a far-red/red ratio greater than 6. Highly shaded leaves in the high plant density treatment accumulated triacylglycerol (TG) up to 25% of total leaf lipid, a 2.4-fold increase in TG on a chlorophyll basis compared to leaves in the upper canopy. Although total polar lipid content was reduced up to 50% in shaded leaves, shade had little affect on the lipid content or composition of thylakoid membranes. Shade did not affect leaf chlorophyll content. Therefore, the changes in leaf lipid composition were not related to senescence. These findings suggest that conditions of low irradiance and/or a high FR/R ratio cause a shift in carbon metabolism toward the accumulation of TG, a storage lipid. Eighteen-carbon fatty acid desaturation was also affected in highly shaded leaves where a reduction in linolenic acid (18:3) content was accompanied by a proportional increase in oleic (18:1) and linoleic (18:2) acids.     

Climate-ameliorating measures influence photosynthetic gas exchange of apple leaves

Sunburn has become one of the major threats to apple fruit production in South Africa and other countries with Mediterranean climate. Some climate‐ameliorating measures have been developed to control sunburn in apples. Effects of the climate‐ameliorating measures, viz. evaporative cooling, Surround^® WP and shade net, on leaf gas exchange of a 5‐year‐old orchard of ‘Cripps’ Pink’ apple were investigated during hot summer days in Stellenbosch, South Africa. Evaporative cooling increased net photosynthetic rate (A) and stomatal conductance (g_s) because of its lowering of leaf temperature and leaf‐to‐air vapour pressure difference (VPD). Shade net also reduced leaf temperature because of reduction in photosynthetic photon flux density (PPFD). Quantum efficiency of photosynthesis was increased under shade net to compensate for reduced PPFD. Shade net also reduced transpiration rate more than A, resulting in increased midday water‐use efficiency. The diurnal trends of A and g_s in the Surround WP and control treatments were similar, indicating limited ameliorative impact of Surround WP. Furthermore, Surround WP typically reduced maximum rate of carboxylation and the light‐saturated rate of electron transport. In all treatments, A decreased by 70% when leaf temperature increased from 35°C to 40°C. In conclusion, all treatments affected leaf photosynthetic gas exchange. Evaporative cooling enhanced leaf A and g_s because of distinct ameliorative effects on leaf temperature and VPD. Shade net reduced leaf temperature with no consistent effects on leaf gas exchange attributes. Surround WP had limited or no impact on leaf temperature and negatively affected leaf gas exchange attributes.     

Leaf area and photosynthesis of newly emerged trifoliolate leaves are regulated by mature leaves in soybean

Leaf anatomy and the stomatal development of developing leaves of plants have been shown to be regulated by the same light environment as that of mature leaves, but no report has yet been written on whether such a long-distance signal from mature leaves regulates the total leaf area of newly emerged leaves. To explore this question, we created an investigation in which we collected data on the leaf area, leaf mass per area (LMA), leaf anatomy, cell size, cell number, gas exchange and soluble sugar content of leaves from three soybean varieties grown under full sunlight (NS), shaded mature leaves (MS) or whole plants grown in shade (WS). Our results show that MS or WS cause a marked decline both in leaf area and LMA in newly developing leaves. Leaf anatomy also showed characteristics of shade leaves with decreased leaf thickness, palisade tissue thickness, sponge tissue thickness, cell size and cell numbers. In addition, in the MS and WS treatments, newly developed leaves exhibited lower net photosynthetic rate (Pn), stomatal conductance (Gs) and transpiration rate (E), but higher carbon dioxide (CO ₂ ) concentration in the intercellular space (Ci) than plants grown in full sunlight. Moreover, soluble sugar content was significantly decreased in newly developed leaves in MS and WS treatments. These results clearly indicate that (1) leaf area, leaf anatomical structure, and photosynthetic function of newly developing leaves are regulated by a systemic irradiance signal from mature leaves; (2) decreased cell size and cell number are the major cause of smaller and thinner leaves in shade; and (3) sugars could possibly act as candidate signal substances to regulate leaf area systemically.     

Photosynthetic activity,chloroplast ultrastructure,and leaf characteristics of high-light and low-light plants and of sun and shade leaves

The photosynthetic CO₂-fixation rates, chlorophyll content, chloroplast ultrastructure and other leaf characteristics (e.g. variable fluorescence, stomata density, soluble carbohydrate content) were studied in a comparative way in sun and shade leaves of beech (Fagus sylvatica) and in high-light and low-light seedlings.

1. Sun leaves of the beech possess a smaller leaf area, higher dry weight, lower water content, higher stomata density, higher chlorophyll a/b ratios and are thicker than the shade leaves. Sun leaves on the average contain more chlorophyll in a leaf area unit; the shade leaf exhibits more chlorophyll on a dry weight basis. Sun leaves show higher rates for dark respiration and a higher light saturation of photosynthetic CO₂-fixation. Above 2000 lux they are more efficient in photosynthetic quantum conversion than the shade leaves.
2. The development of HL-radish plants proceeds much faster than that of LL-plants. The cotyledons of HL-plants show a higher dry weight, lower water content, a higher ratio of chlorophyll a/b and a higher gross photosynthesis rate than the cotyledons of the LL-plants, which possess a higher chlorophyll content per dry weight basis. The large area of the HL-cotyledon on the one hand, as well as the higher stomata density and the higher respiration rate in the LL-cotyledon on the other hand, are not in agreement with the characteristics of sun and shade leaves respectively.
3. The development, growth and wilting of wheat leaves and the appearance of the following leaves (leaf succession) is much faster at high quanta fluence rates than in weak light. The chlorophyll content is higher in the HL-leaf per unit leaf area and in the LL-leaf per g dry weight. There are no differences in the stomata density and leaf area between the HL- and LL-leaf. There are fewer differences between HL- and LL-leaves than in beech or radish leaves.
4. The chloroplast ultrastructure of shade-type chloroplasts (shade leaves, LL-leaves) is not only characterized by a much higher number of thylakoids per granum and a higher stacking degree of thylakoids, but also by broader grana than in sun-type chloroplasts (sun leaves, HL-leaves). The chloroplasts of sun leaves and of HL-leaves exhibit large starch grains.
5. Shade leaves and LL-leaves exhibit a higher maximum chlorophyll fluorescence and it takes more time for the fluorescence to decline to the steady state than in sun and HL-leaves. The variable fluorescence VF (ratio of fluorescence decrease to steady state fluorescence) is always higher in the sun and HL-leaf of the same physiological stage (maximum chlorophyll content of the leaf) than in the shade and LL-leaf. The fluorescence emission spectra of sun and HL-leaves show a higher proportion of chlorophyli fluorescence in the second emission maximum F₂ than shade and LL-leaves.
6. The level of soluble carbohydrates (reducing sugars) is significantly higher in sun and HL-leaves than in shade and LL-leaves and even reflects changes in the amounts of the daily incident light.
7. Some but not all characteristics of mature sun and shade leaves are found in HL- and LL-leaves of seedlings. Leaf thickness, dry weight, chlorophyll content, soluble carbohydrate level, photosynthetic CO₂-fixation, height and width of grana stacks and starch content, are good parameters to describe the differences between LL- and HL-leaves; with some reservations concerning age and physiological stage of leaf, a/b ratios, chlorophyll content per leaf area unit and the variable fluorescence are also suitable.