Changes of Starch and Sorbitol in Leaves Before and After Removal of Fruits from Peach Trees

Changes in contents of nonstructural carbohydrates in leaves,as well as some characteristics of leaves before and after fruitremoval, were investigated in potted peach (Prunus persica L.)trees. Leaf area and dry mass per unit leaf area (SLW) at thefruit-maturation stage decreased with increasing numbers ofpeaches per tree, whereas the chlorophyll content per unit areain leaves of fruiting trees increased. The chlorophyll contentdecreased more rapidly upon removal of fruit than that in leavesof fruiting trees. The starch content per unit dry mass in leavesof fruiting trees at the fruit-maturation stage was lower thanthat in leaves of non-fruiting trees. Starch accumulated significantlyin leaves within 1 d of removal of fruit during the fruit-maturationstage and continued to increase thereafter. The accumulationof starch after removal of fruit occurred more rapidly thanthe decrease in chlorophyll content. Reducing and non-reducingsugars (total sugars) per unit dry mass in the leaves were higherin fruiting trees than in non-fruiting trees. After fruit removal,the total sugar content of leaves increased temporarily andthen gradually decreased. The sorbitol content per unit freshmass in leaves of fruiting trees during the fruit-maturationstage was slightly higher than that in leaves of non-fruitingtrees. One day after removal of fruit, the sorbitol contentincreased in parallel with the accumulation of starch and remainedhigh. The sucrose content of leaves did not change markedlyupon removal of fruit. Prunus persica L.; peach leaves; nonstructural carbohydrate; starch and sorbitol; fruit removal     

Relationships between Carbon Assimilation, Partitioning, and Export in Leaves of Two Soybean Cultivars

To evaluate leaf carbon balance during rapid pod-fill in soybean (Glycine max [L.] Merrill), measurements were made of CO₂ assimilation at mid-day and changes in specific leaf weight, starch, and sucrose concentrations over a 9-hour interval. Assimilate export was estimated from CO₂ assimilation and leaf dry matter accumulation. Chamber-grown `Amsoy 71' and `Wells' plants were subjected on the day of the measurements to one of six photosynthetic photon flux densities in order to vary CO₂ assimilation rates.

Rate of accumulation of leaf dry matter and rate of export both increased as CO₂ assimilation rate increased in each cultivar.

Starch concentrations were greater in Amsoy 71 than in Wells at all CO₂ assimilation rates. At low CO₂ assimilation rates, export rates in Amsoy 71 were maintained in excess of 1.0 milligram CH₂O per square decimeter leaf area per hour at the expense of leaf reserves. In Wells, however, export rate continued to decline with decreasing CO₂ assimilation rate. The low leaf starch concentration in Wells at low CO₂ assimilation rates may have limited export by limiting carbon from starch remobilization.

Both cultivars exhibited positive correlations between CO₂ assimilation rate and sucrose concentration, and between sucrose concentration and export rate. Carbon fixation and carbon partitioning both influenced export rate via effects on sucrose concentration.

     

Energy requirement for foliage formation is not constant along canopy light gradients in temperate deciduous trees

Foliage construction cost (glucose requirement for formation of a unit foliar biomass, G , kg glu kg⁻¹), chemical composition and morphology were examined along a light gradient across the canopies in five deciduous species, which ranked according to increasing shade-tolerance as Populus tremula < Fraxinus excelsior < Tilia cordata = Corylus avellana < Fagus sylvatica . Light conditions in the canopy were estimated by a hemispheric photographic technique, allowing ranking of sample locations according to long-term light input incident to the sampled leaves (relative irradiance). G and foliage carbon concentration increased with increasing relative irradiance in F. excelsior , T. cordata and C. avellana , but wereindependent of irradiance in F. sylvatica and P. tremula . However, if G of non-structural-carbohydrate-free dry mass was considered, it also increased with increasing relative irradiance in P. tremula . A positive correlation between the concentration of carbon-rich lignin and irradiance, probably a result of the acclimation to greater water stress at higher light, was the major reason for the light-dependence of G . Lignin concentrations were highest in more shade-tolerant species, resulting in greatest carbon concentrations in these species. Since carbon concentration and G are directly linked, the leaves of shade-tolerant species were also more expensive to construct. As the result of these effects, G increased faster with increasing leaf dry mass per area which was mainly determined by relative irradiance, in shade-tolerators. Given that shade-tolerant species had lower leaf dry mass per area at common irradiance and that this saturated at lower relative irradiance than leaf dry mass per area in the intolerant species, it was concluded that enhanced energy requirements for foliage construction might constrain species morphological plasticity and the upper limit of leaf dry mass per area attainable at high light.     

Root Respiration and Growth in Plantago major as Affected by Vesicular-Arbuscular Mycorrhizal Infection

Effects of vesicular-arbuscular mycorrhizal (VAM) infection and P on root respiration and dry matter allocation were studied in Plantago major L. ssp. pleiosperma (Pilger). By applying P, the relative growth rate of non-VAM controls and plants colonized by Glomus fasciculatum (Thaxt. sensu Gerdemann) Gerdemann and Trappe was increased to a similar extent (55-67%). However, leaf area ratio was increased more and net assimilation rate per unit leaf area was increased less by VAM infection than by P addition. The lower net assimilation rate could be related to a 20 to 30% higher root respiration rate per unit leaf area of VAM plants. Root respiration per unit dry matter and specific net uptake rates of N and P were increased more by VAM infection than by P addition. Neither the contribution of the alternative respiratory path nor the relative growth rate could account for the differences in root respiration rate between VAM and non-VAM plants. It was estimated that increased fungal respiration (87%) and ion uptake rate (13%) contributed to the higher respiratory activity of VAM roots of P. major.     

Morphological and chemical leaf composition of Mediterranean evergreen tree species according to leaf age

Changes in morphology [leaf dry mass per unit area (LMA), thickness and density] and chemical composition (macronutrients and fibres content) in different age leaves of eight evergreen Mediterranean woody species were investigated. LMA and leaf thickness increased with leaf age increasing. Young tissues possessed higher concentrations of N, P, K, and Mg and lower Ca concentrations on a dry mass basis. However, mineral content was independent of age on leaf area basis (except for Ca content) suggesting that the changes in mineral concentration with leaf ageing are due to dilution in the larger dry mass accumulated in the oldest leaves. Leaf tissue density (LTD) increased during the first year of the leaf life. Lignin and hemicellulose concentrations did not vary along leaf life and the cellulose concentration increased with leaf age in most species between the current-year and the one-year old leaves. Our results suggested that physical leaf reinforcement with a higher cellulose concentration and density might be a leaf response to the unfavourable climatic conditions during the first winter.     

Effects of Nitrate Application on Amaranthus powellii Wats. : I. Changes in Photosynthesis, Growth Rates, and Leaf Area

Physiological effects of different nitrate applications were studied using the C₄ plant, Amaranthus powellii Wats. Plants were grown in a controlled environment chamber and watered daily with nutrient solutions containing 45, 10, 5, or 1 millimolar nitrate. Chloride and sulfate were used to keep the cation and phosphate concentrations constant. Total leaf nitrogen concentration, chlorophyll concentration, specific leaf mass, leaf area, relative growth rate, relative leaf growth rate, unit leaf rate (increase of dry mass per unit leaf area per day), net photosynthetic rate, and incident quantum yield decreased with decreasing nitrate concentration. The per cent decrease of unit leaf rate was similar to the decrease of light-saturated net photosynthetic rate; however, the decrease in relative growth rate was less than that of unit leaf rate because leaf area ratio (leaf area per unit dry mass) increased with decreasing nitrate concentration. Essential mineral concentrations per unit leaf area were about equal among all treatments. Leaf expansion, determined by stomatal density, decreased except for the 1 millimolar treatment which showed relatively more cell expansion but less cell division. Decreased nitrate application was correlated with higher osmotic potentials and lower pressure potentials (determined by pressure-volume curves), whereas leaf water potentials were equal among treatments. Even though total leaf area and shoot mass decreased with decreasing applied nitrate, the increase of the leaf area ratio may be related to selection for the highest possible growth rate.     

Leaf Phosphate Status, Photosynthesis, and Carbon Partitioning in Sugar Beet: III. Diurnal Changes in Carbon Partitioning and Carbon Export

The effect of low phosphate supply (low P) was determined on the diurnal changes in the rate of carbon export, and on the contents of starch, sucrose, glucose, and fructose 2,6-bisphosphate (F2,6BP) in leaves. Low-P effects on the activities of a number of enzymes involved in starch and sucrose metabolism were also measured. Sugar beets (Beta vulgaris L. cv. F58-554H1) were cultured hydroponically in growth chambers and the low-P treatment induced nutritionally. Low-P treatment decreased carbon export from the leaf much more than it decreased photosynthesis. At growth chamber photon flux density, low P decreased carbon export by 34% in light; in darkness, export rates fell but more so in the control so that the average rate in darkness was higher in low-P leaves. Low P increased starch, sucrose, and glucose contents per leaf area, and decreased F2, 6BP. The total extractable activities of enzymes involved in starch and sucrose synthesis were increased markedly by low P, e.g. adenosine 5-diphosphoglucose pyrophosphorylase, cytoplasmic fructose-1,6-bisphosphatase, uridine 5-diphosphoglucose pyrophosphorylase, and sucrose-phosphate synthase. The activities of some enzymes involved in starch and sucrose breakdown were also increased by low P. We propose that plants adapt to low-P environments by increasing the total activities of several phosphatases and by increasing the concentrations of phosphate-free carbon compounds at the expense of sugar phosphates, thereby conserving Pi. The partitioning of carbon among the various carbon pools in low-P adapted leaves appears to be determined in part by the relative capacities of the enzymes for starch and sucrose metabolism.     

Effects of Water Stress on Photosynthesis and Carbon Partitioning in Soybean (Glycine max [L.] Merr.) Plants Grown in the Field at Different CO(2) Levels

The effects of water stress and CO₂ enrichment on photosynthesis, assimilate export, and sucrose-P synthase activity were examined in field grown soybean plants. In general, leaves of plants grown in CO₂-enriched atmospheres (300 microliters per liter above unenriched control, which was 349 ± 12 microliters per liter between 0500 and 1900 hours EST over the entire season) had higher carbon exchange rates (CER) compared to plants grown at ambient CO₂, but similar rates of export and similar activities of sucrose-P synthase. On most sample dates, essentially all of the extra carbon fixed as a result of CO₂ enrichment was partitioned into starch. CO₂-enriched plants had lower transpiration rates and therefore had a higher water use efficiency (milligrams CO₂ fixed per gram H₂O transpired) per unit leaf area compared to nonenriched plants. Water stress reduced CER in nonenriched plants to a greater extent than in CO₂-enriched plants. As CER declined, stomatal resistance increased, but this was not the primary cause of the decrease in assimilation because internal CO₂ concentration remained relatively constant. Export of assimilates was less affected by water stress than was CER. When CERs were low as a result of the imposed stress, export was supported by mobilization of reserves (mainly starch). Export rate and leaf sucrose concentration were related in a curvilinear manner. When sucrose concentration was above about 12 milligrams per square decimeter, obtained with nonstressed plants at high CO₂, there was no significant increase in export rate. Assimilate export rate was also correlated positively with SPS activity and the quantitative relationship varied with CER. Thus, export rate was a function of both CER and carbon partitioning.     

Abscisic acid mimics effects of dehydration on area expansion and photosynthetic partitioning in young soybean leaves

Abstract. Leaf area expansion, photosynthetic carbon dioxide uptake and leaf dry mass accumulation were compared for expanding leaves of well-watered soybean ( Glycine max [L.] Merr.) plants, mildly dehydrated plants and well-watered plants treated with abscisic acid (ABA). Both ABA treatment and dehydration reduced area expansion in the light and over a 24 h period without decreasing the photosynthetic rates of expanding leaves. Dry mass accumulation during the light was less in ABA-treated and water-stressed leaves than in control leaves, with no differences among treatments in leaf mass per unit of area. ABA treatment and water stress both increased export of carbon from expanding leaves in the light. ABA applied near the end of the light period also increased export of carbon during the following dark period. However, it is unlikely that decreased availability of photosynthate caused slow expansion in the ABA and dehydration treatments, because expansion rates were not slowed in plants kept in dim light, even though photosynthetic rates and dry mass accumulation rates were greatly reduced. The data suggest that ABA may mediate the effects of mild dehydration on leaf area expansion and partitioning of photosynthate.     

Carbon partitioning and export from mature cotton leaves

The partitioning of carbon in intact, mature cotton (Gossypium hirsutum L.) leaves was examined by steady-state ¹⁴CO₂ labeling. Plants were exposed to dark periods of varying lengths, followed by similar illuminated labeling periods. These treatments produced leaves with a range of starch and soluble sugar contents, carbon exchange, and carbon export rates. Export during the illuminated periods was neither highly correlated with photosynthesis nor was export during the illuminated periods significantly different among the treatments. In contrast, the rate of subsequent nocturnal carbon export from these leaves varied widely and was found to be highly correlated with leaf starch content at the end of the illumination period (r = 0.934) and with nocturnal leaf respiration (r = 0.954). Leaves which had accumulated the highest levels of starch (about 275 micrograms per square centimeter) by the end of the illumination period exhibited nocturnal export rates very similar to those during the daylight hours. Leaves which accumulated starch to only 50 to 75 micrograms per square centimeter virtually ceased nocturnal carbon export. For leaves with starch accumulations of between 50 and 275 micrograms per square centimeter, nocturnal export was directly proportional to leaf starch at the end of the illumination period. After the nocturnal export rate was established, it continued at a constant rate throughout the night even though leaf starch and sucrose contents declined.     

Diel patterns of leaf C export and of main shoot growth for Flaveria linearis with altered leaf sucrose-starch partitioning

Diel C export from source leaves of two Flaveria linearis lines [85-1: high cytosolic fructose-1,6-bisphosphatase (cytFBPase) and 84-9: low cytFBPase] were estimated using three methods, including leaf steady-state (14)CO(2) labelling, leaf metabolite analysis, and leaf dry mass analysis in conjunction with leaf CO(2) exchange measurements. Synthesis and accumulation of starch during the daytime were much higher in 84-9. Relative (14)C-export (export as a % of photosynthesis) in the light was 36% higher in 85-1. The diel export patterns from (14)C-analyses correlated with those based on metabolite or dry weight/gas exchange analyses during the daytime, but not during the night. Night-time export estimated from (14)C-disappearance was 3.6 times lower than those estimated using the other methods. Even though the starch degradation at night was greater for 84-9, night-time export in 84-9 was similar to 85-1, since 84-9 showed both higher respiration and accumulation of soluble sugars (i.e. glucose) at night. Patterns of (14)C allocation to sink organs were also different in the two lines. Main stem growth was less in 84-9, being reduced most in the light when leaf export was lower relative to 85-1. Supplementation with sucrose for 1 h daily via the roots at a time when leaf export in 84-9 was low relative to 85-1 increased the stem growth rate of 84-9 to a level similar with that of 85-1. This study provides evidence that diel C availability predicted by source strength (e.g. C-export rate) influences main stem extension growth and the pattern of sink development in F. linearis.     

Effect of Water Stress on Photosynthesis, Leaf Characteristics and Productivity of Field-Grown Nicotiana tabacum L. Genotypes Selected for Survival at Low CO2

Dry mass production, leaf characteristics and diurnal photosynthesisof two N. tabacum L. genotypes selected for survival at lowCO₂ and the parent Wisconsin-38 (control plants) were measuredon water-stressed and well-watered plants in the field. Differencesin photosynthesis per unit leaf area were small and not significantbetween genotypes, but different patterns of photosynthesiswere observed in stressed and non-stressed plants, with waterstress reducing total net carbon fixation by 45% in all genotypes.More dry mass was produced by the selected genotypes than byWisconsin-38 under irrigation. Production was smaller and thesame for all three genotypes when stressed. The increased drymass of the selected genotypes was related to greater totalleaf area per plant which was accompanied by more cells perunit leaf area but smaller cell volume than in the control plants.The decrease in dry mass production under water stress was relatedto a decrease in total leaf area per plant and a decrease incell number per unit leaf area; however, cell volume increased. Key words: Water stress, photosynthesis, productivity, leaf cells, genotype     

Glyphosate inhibits photosynthesis and allocation of carbon to starch in sugar beet leaves

Application of glyphosate (N-[phosphonomethyl] glycine) to exporting leaves of sugar beet (Beta vulgaris, L.) during the day lowered stomatal conductance and carbon fixation. Allocation of newly fixed carbon to foliar starch accumulation was nearly completely inhibited, being decreased by the same amount as net carbon fixation. In contrast, decreasing net carbon fixation in untreated leaves by lowering CO₂ concentration caused starch accumulation to decrease, but only in the same proportion as net carbon fixation. Shikimate level increased 50-fold in treated leaves but the elevated rate of carbon accumulation in shikimate was only 4% of the decrease in the rate of starch accumulation. Application of steady state labeling with ¹⁴CO₂ to exporting leaves confirmed the above changes in carbon metabolism, but revealed no other major daytime differences in the ¹⁴C-content of amino acids or other compounds between treated and control leaves. Less ¹⁴C accumulated in treated leaves because of decreased fixation, not increased export. The proportion of newly fixed carbon allocated to sucrose increased, maintaining export at the level in control leaves. Returning net carbon exchange to the rate before treatment restored starch accumulation fully and prevented a decrease in export during the subsequent dark period.     

Relation of increased potassium nutrition to photosynthesis and translocation of carbon

Effects of supplying K⁺ at 2 or 10 millimolarity concentration on net carbon exchange and translocation of products of photosynthesis were studied in plants of Beta vulgaris L. (var. Klein E). Transport of K⁺ into and out of leaves was studied with ⁴²K over a 3-day period. Increasing the K⁺ supplied to the roots from 2 millimolarity, a level just sufficient to overcome obvious deficiency symptoms, to 10 millimolarity resulted in a gradual accumulation of K⁺ per unit area and an increased export of K⁺ to sink regions. No significant increase in net carbon exchange was observed in leaves that had accumulated a high level of K⁺ per unit area. Initiation rate, total area, and total fresh weight of leaves of plants with K⁺ supplied at 10 millimolarity was similar to that for leaves from plants at a 2 millimolarity level. Shoot/root ratio and dry weight accumulation, which are indicative of translocation and partitioning over the long term, were independent of K⁺ supply in the 2 to 10 millimolarity range. Accumulation of K⁺ by exporting leaves and its subsequent recirculation to sinks increased when K⁺ supply was increased in this range but did not appear to affect carbon nutrition even after a long period.     

The Glycine-Glomus-Rhizobium Symbiosis : VII. Photosynthetic Nutrient-Use Efficiency in Nodulated, Mycorrhizal Soybeans

Four consecutive trifoliate leaves of 56-day-old symbiotic or nonsymbiotic soybean plants were evaluated individually for CO₂ exchange rates (CER), leaf area and dry weight, and leaf N, P, and starch concentrations. Plants had been inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus mosseae and Rhizobium japonicum, with either of the endophytes alone, or with neither at time of planting. Plants lacking one or both endophytes received N and/or P fertilizers to produce plants of equal total leaf dry weight in all four treatments. Photosynthetic P-use efficiency (CER per unit leaf P) was higher in the leaves of VAM plants than in P-fertilized plants regardless of the N source (N₂ fixation or combined N). Photosynthetic N-use efficiency was also higher in VAM than in non-VAM plants, but it was affected by the N source, with higher CER in the nodulated plants. The greatest differences in CER, starch accumulation and leaf area were found between the nonsymbiotic plants and those with both endophytes. Statistical evaluations of leaf parameters for treatment or nutrient concentration (N and P) effects between the tri-partite and the nonsymbiotic treatments showed significant changes in concentration of P, but not N, with decreasing leaf age. Both endophytes apparently enhance CO₂ fixation at N and/or P concentrations lower than those of the nonsymbiotic plants. The effects of the endophytes on CO₂ fixation were additive.     

The relationship between leaf composition and morphology at elevated CO2 concentrations

The composition and morphology of leaves exposed to elevated [CO₂] usually change so that the leaf nitrogen (N) per unit dry mass decreases and the leaf dry mass per unit area increases. However, at ambient [CO₂], leaves with a high leaf dry mass per unit area usually have low leaf N per unit dry mass. Whether the changes in leaf properties induced by elevated [CO₂] follow the same overall pattern as that at ambient [CO₂] has not previously been addressed. Here we address this issue by using leaf measurements made at ambient [CO₂] to develop an empirical model of the composition and morphology of leaves. Predictions from that model are then compared with a global database of leaf measurements made at ambient [CO₂]. Those predictions are also compared with measurements showing the impact of elevated [CO₂]. In the empirical model both the leaf dry mass and liquid mass per unit area are positively correlated with leaf thickness, whereas the mass of C per unit dry mass and the mass of N per unit liquid mass are constant. Consequently, both the N:C ratio and the surface area:volume ratio of leaves are positively correlated with the liquid content. Predictions from that model were consistent with measurements of leaf properties made at ambient [CO₂] from around the world. The changes induced by elevated [CO₂] follow the same overall trajectory. It is concluded that elevated [CO₂] enhances the rate at which dry matter is accumulated but the overall trajectory of leaf development is conserved.     

Increased chilling tolerance and altered carbon metabolism in tomato leaves following application of mechanical stress

We investigated the effects of brushing on the chilling tolerance and metabolism of nonstructural carbohydrates (soluble sugars and starch) in tomato leaves before, during and after a chilling stress. Tomato plants ( Lycopersicon esculentum Mill. cv. Caruso) were cultivated either without mechanical stress application (control plants) or with daily brushing treatments for 15 days (brushed plants), prior to a 7-day chilling treatment (8/5°C day/night). Brushing resulted in shorter plants with a 34% reduction in leaf dry weight per area and a 59% reduction of soluble sugars and starch, on a dry weight basis. The sugar to starch ratio was not affected by brushing. A greater chilling tolerance in the brushed plants was demonstrated by the maintenance of a significantly higher PSII efficiency in brushed plants (42%) compared to that of the control plants (30%) after 7 days of chilling treatment, less visible damage to the leaf tissue, and a more rapid resumption of growth during 3 days of recovery as compared to control plants. During the chilling treatment levels of soluble sugars per leaf dry weight increased 15-fold in the brushed plants and 5-fold in control plants. In the present study we have demonstrated that brushing can increase chilling tolerance in tomato plants. The observed differences in chilling tolerance and concentration of soluble sugars in the leaves may indicate an involvement of soluble sugar levels in acclimation to chilling.     

Growth and carbon partitioning in compatible and incompatible peach/plum grafts

The shoot growth of compatible ( Prunus persica L. Batsch cv. Springtime grafted on Prunus cerasifera L. Ehrh cv. Myrobolan P2032) and incompatible ( Prunus persica L. Batsch cv. Springtime grafted on Prunus cerasifera L. Ehrh cv. Myrobolan P18) peach/plum grafts was observed over a period of 100 days after grafting under controlled conditions. Leaf and root activities were determined by studying carbon assimilation and partitioning, leaf mineral contents and water relations. Shoot length and leaf number were not significantly affected in the incompatible combination during the first 55 days after grafting, but then, shoot growth rate was significantly reduced. Final total dry weights of the shoot were similar in both graft combinations. The incompatible combination did not show any water stress. Soluble sugar and starch contents increased in the leaves of the incompatible combination, accounting for about 36% of the increase of leaf dry weight per unit area. Photosynthesis was affected by the compatibility of the grafts. Leaf nitrogen content (% dry weight) fell in the incompatible graft combination 65 days after grafting. However, nitrogen content on an area basis was not affected. The possibility of nitrogen stress is discussed.     

Carbon Partitioning and Assimilation as Affected by Nitrogen Deficiency in Cassava

Plants of cassava (Manihot esculenta Crantz) were raised in a sand root medium watered with nutrient solutions, under greenhouse conditions. As the N-supply increased, shoot dry mass was enhanced to a greater extent than root dry mass, thus leading to an increased shoot to root ratio. In leaves, contents of total soluble saccharides, non-reducing saccharides, and inorganic phosphate increased linearly with increasing N-supply. An opposite response was found for reducing saccharides and starch. In general, content of non-reducing saccharides was considerably greater than starch content. Activity of sucrose synthase was not detected, regardless of the N-treatments; by contrast, activity of neutral and acid invertases increased with increasing N-availability. Roots accumulated more total soluble saccharides, but less reducing saccharides and starch, as the N-supply increased. Photosynthetic rates decreased with increasing N-deficiency. Such a decrease was circumstantially associated to reducing saccharide, but not starch, accumulation. Results suggest a limited capacity for carbon export from source leaves under N-limitation. This revised version was published online in August 2006 with corrections to the Cover Date.     

The analysis of the causes of variability of the relationship between leaf dry mass and area in plants

The lamina dry mass: area ratio (LMA - Leaf Mass per Area) is a quite variable trait. Leaf dry mass consists of symplast mass (a set of all leaf protoplasts) and apoplast mass (a set of all cell walls in a leaf). The ratio between symplast and apoplast masses is positively related to any functional trait of leaf calculated per unit of dry mass. The value of this ratio is defined by cells size and their number per unit of leaf area, number of mesophyll cells layers and their differentiation between palisade and spongy ones, and also by density of cells packing. The LMA value is defined by leaf thickness and density. The extent and direction of variability in both leaf traits define the extent and direction of variability in LMA. Negative correlation between leaf thickness and density reduces the level of LMA variability. As a consequence of this correlation the following pattern emerges: the thinner a leaf, the denser it is. Changes in the traits that define the LMA value take place both within a species under the influence of environmental factors and between species that differ in leaf structure and functions. Light is the most powerful environmental factor that influences the LMA, increase in illumination leading to increase in LMA. This effect occurs during leaf growth at the expense of structural changes associated with the reduction of symplast/apoplast mass ratio. Under conditions of intense illumination, LMA may increase due to accumulation of starch. With regard to the majority of leaf functions, the mass of starch may be ascribed to apoplast. Starch accumulation in leaves is observed also under conditions of elevated CO2 concentration in the air. Under high illumination, however, LMA increases also due to increased apoplast contribution to leaf dry mass. Scarce mineral nutrition leads to LMA increase due to lowering of growth zones demands for phothosyntates and, therefore, to increase in starch content of leaves. High level of mineral nutrition during leaf growth period leads to LMA increase at the expense of mesophyll thickening where components of photosynthesis system are located. When additional environmental factors are involved, starch accumulation may be partly responsible for increase in LMA. LMA increase at the expense of starch accumulation, unlike that at the expense of mesophyll thickening, is accompanied by increased leaf density. Under conditions of water deficiency LMA increases, which in mature leaf may be caused by starch accumulation. LMA increase during leaf growth period under conditions of water deficiency is associated with decrease in the symplast/apoplast mass ratio.