Distribution and redistribution of sulphur taken up from nutrient solution during vegetative growth in barley

Barley plants were grown in a nutrient solution containing 25 μ M sulphate and the roots were pulsed with [³⁵S]sulphate for 48-h periods at 6 different times between the emergence of leaf 5 (L5) and the emergence of leaf 8 (L8). Growth was continued in unlabelled solution until the emergence of L10. Within the shoot system sulphur was directed principally into the leaf undergoing expansion. A large proportion of the ³⁵S-label delivered to young expanding leaves (> 40% of full expansion) did not occur at the time of the pulse, but subsequently during the ensuing chase indicating slow redistribution of sulphur from another site. During the later stages of leaf expansion (40–100%), most of the sulphur entered the leaf during the pulse, suggesting that sulphur was delivered more directly from the nutrient solution. Up to 75% of the sulphur delivered to L3–L6 at the time they approached or attained full expansion (70–100%) was re-exported. At least some of the sulphur exported from fully expanded leaves was redistributed to developing leaves.     

Effect of Sulfur Nutrition on the Redistribution of Sulfur in Vegetative Soybean Plants

Soybean (Glycine max L.) plants were grown with sulfate at 2 (S2) or 20 [mu]M (S20) and treated with [35S]sulfate between d 36 and 38. Growth was continued with or without 20 [mu]M sulfate (i.e. S2 -> S0, S2 -> S20, etc.). When the leaves of S20 -> S20 plants were 70% expanded, they exported S and 35S label from the soluble fraction, largely as sulfate, to new expanding leaves. However, 35S label in the insoluble fraction was not remobilized. Very little of the 35S label in the soluble fraction of the leaves of S20 -> S0 plants was redistributed; most was incorporated into the insoluble fraction. The low levels of S remobilization from the insoluble fraction were attributed to the high level of N in the nutrient solution (15 mM). Most of the 35S label in S2 plants at d 38 occurred in the soluble fraction of the roots. In S2 -> S0 plants the 35S label was incorporated into the insoluble fraction of the roots, but in S2 -> S20 plants 35S label was rapidly exported to leaves 3 to 6. It was concluded that the soluble fraction of roots contains a small metabolically active pool of S and another larger pool that is in slow equilibrium with the small pool.     

Response of cassava leaf area expansion to water deficit: cell proliferation, cell expansion and delayed development

BACKGROUND AND AIMS: Cassava (Manihot esculenta) is an important food crop in the tropics that has a high growth rate in optimal conditions, but also performs well in drought-prone climates. The objectives of this work were to determine the effects of water deficit and rewatering on the rate of expansion of leaves at different developmental stages and to evaluate the extent to which decreases in cell proliferation, expansion, and delay in development are responsible for reduced growth. METHODS: Glasshouse-grown cassava plants were subjected to 8 d of water deficit followed by rewatering. Leaves at 15 developmental stages from nearly full size to meristematic were sampled, and epidermal cell size and number were measured on leaves at four developmental stages. KEY RESULTS: Leaf expansion and development were nearly halted during stress but resumed vigorously after rewatering. In advanced-stage leaves (Group 1) in which development was solely by cell expansion, expansion resumed after rewatering, but not sufficiently for cell size to equal that of controls at maturity. In Group 2 (cell proliferation), relative expansion rate and cell proliferation were delayed until rewatering, but then recovered partially, so that loss of leaf area was due to decreased cell numbers per leaf. In Group 3 (early meristematic development) final leaf area was not affected by stress, but development was delayed by 4-6 d. On a plant basis, the proportion of loss of leaf area over 26 d attributed to leaves at each developmental stage was 29, 50 and 21 % in Group 1, 2 and 3, respectively. CONCLUSIONS: Although cell growth processes were sensitive to mild water deficit, they recovered to a large extent, and much of the reduction in leaf area was caused by developmental delay and a reduction in cell division in the youngest, meristematic leaves.     

Phloem transport of sulfur in Ricinus

Mature leaves of Ricinus communis fed with ³⁵SO ₄ ^2- in the light export labeled sulfate and reduced sulfur compounds by phloem transport. Only 1–2% of the absorbed radiosulfur is exported to the stem within 2–3 h, roughly 12% of ³⁵S recovered was in reduced form. The composition of phloem translocate moving down the stem toward the root was determined from phloem exudate: 20–40% of the ³⁵S moved in the form of organic sulfur compounds, however, the bulk of sulfur was transported as inorganic sulfate. The most important organic sulfur compound translocated was glutathione, carrying about 70% of the label present in the organic fraction. In addition, methionine and cysteine were involved in phloem sulfur transport and accounted for roughly 10%. Primarily, the reduced forms of both, glutathione and cysteine are prsent in the siever tubes.Abbreviations CySH cysteine - GSH glutathione - GSSG glutathione disulfide - NEM N-ethylmaleimide - CyS-SCy cystine     

Inhibition of sulphur redistribution into new leaves of vegetative soybean by excision of the maturing leaf

When soybean plants are pulsed with [³⁵S]sulphate, label is subsequently redistributed from the roots to the leaves. This confounds studies to measure the redistribution of label from leaves. Accordingly, soybean plants ( Glycine max [L.] Merr. cv. Stephens) were grown in 20 μ M sulphate and a small portion of the root system (donor root) was pulsed with [³⁵S]sulphate for 24 h. After removing the donor root, the plants were transferred into unlabelled solution, either without sulphate (S20→SO) or with 20 μ M sulphate (S20→20) (intact plants). Also at this time, the expanding leaf (L3) was excised from half of the plants in each treatment (excised plants). Immediately after the pulse, only ca 15% of the label occurred in the roots and ca 40% in the expanding leaf, L3, mostly in the soluble fraction. In intact S20→20 plants, ³⁵S-label was exported from the soluble fraction of L3, mostly as sulphate, whilst L4 and L5 imported label. Similar responses occurred in S20→SO plants except that export of label from L3 was more rapid. Excision of L3 from S20→S20 plants inhibited labelling of leaves L4-L6 but not total sulphur, whereas in S20→SO plants, excision of L3 inhibited the import of both total sulphur and ³⁵S-label in leaves L4, L5 and L6. The data suggest that the soluble fraction of almost fully expanded leaves is an important reserve of sulphur for redistribution to growing leaves. The ³⁵S-label in the root system exhibited fluctuations consistent with its proposed role in the recycling of soluble sulphur from the leaves.     

Contrasting leaf development within the genus Syzgium

Leaf developmental patterns were characterized in four rainforest tree species of Syzgium. Leaf optical properties, pigment changes, expansion characteristics, stomatal development, and photosynthetic rates were studied. In both S. luehmannii and S. wilsonii photosynthetic development was delayed until after full leaf expansion. Rates of O2 evolution were negative during expansion of S. luehmannii and S. wilsonii leaves and stomatal conductance was 10-20 mmol m^-2 s^-1 lower than for corresponding leaves of S. moorei. Stomatal conductance showed that the development of functional stomata was delayed until after full leaf expansion in S. luehmannii and S. wilsonii, however, low stomatal conductance was not responsible for the lack of photosynthetic potential during leaf expansion in these species. Leaves of S. luehmannii and S. wilsonii required less than 10 d for full leaf expansion and contained anthocyanin during expansion. In contrast, leaves of S. moorei and S. corynanthum expanded slowly (20-40 d required for full leaf expansion), exhibited positive rates of O2 evolution and did not accumulate anthocyanin. In S. luehmannii and S. wilsonii anthocyanin was located in the vacuole of distinct cell layers just below the upper epidermis and the possible functions of anthocyanin accumulation are discussed. This is the first report where such variation in leaf development has been characterized in the one genus.Key words: Anthocyanin, leaf expansion, photosynthetic development, delayed leaf greening, stomatal development.      

常绿阔叶林中栲树展叶期叶片的虫食格局

叶片的虫食主要发生在展叶期。虽然展叶期短暂, 它却可能是了解植食性昆虫和植物之间相互关系的关键。为了解栲树(Castanopsis fargesii)在展叶期叶片的虫食格局和展叶方式对叶片虫食的影响, 研究了栲树展叶期内的虫食动态变化, 结果表明: 栲树展叶的两个阶段(折叠期和打开期), 虫食叶片的格局存在较大的差异, 打开阶段的日虫食频度和日虫食率显著高于折叠阶段(F_{1, 32}=8.97, p=0.005 4; F_{1, 32}=12.38, p=0.001 4), 展叶期最终叶片虫食频度为50.72%, 叶片虫食率为8.25%。折叠期叶片主要受到低强度的虫食, 打开期叶片虫食则以较大强度的虫食为主。展叶期叶片的虫食主要发生在夜间, 夜间虫食率显著高于日间虫食率(t=2.51, p=0.017), 变化趋势与日虫食率一致。栲树叶片在展叶的两个阶段可能采用了不同的防御对策。     

Metabolization of elemental sulfur in wheat leaves consecutive to its foliar application

The qualitative and quantitative aspects of elemental sulfur metabolization in wheat leaves and its effect upon photosynthetic metabolism were studied through the application of micronized sulfur upon the third leaf. Energy-dispersive x-ray analysis combined with scanning electron microscopy emphasized the existence of a sulfur peak associated with a strong potassium peak in the spectra of different tissue regions for treated leaves only, supplying an original evidence of sulfur uptake. Experiments with³⁵S-labeled micronized sulfur showed that about 2% of the labeled S was absorbed and metabolized into cystine, methionine, glutathione, and sulfate. The close correlation between the excess of oxygen uptake and oxygen needs for sulfur oxidation in conjunction with the absence of hydrogen sulfide released by treated leaves support direct and fast oxidation of sulfur into sulfate according to a pathway still unclear but independent of photosynthetic CO₂ metabolism in treated leaf. The mechanisms involved in the primary metabolism of element sulfur in wheat therefore appear to be different from those in fungi.     

Enhanced antioxidant protection at the early stages of leaf expansion in ginkgo under natural environmental conditions

Photosynthetic pigments, gas exchange, chlorophyll (Chl) a fluorescence kinetics, antioxidant enzymes and chloroplast ultrastructure were investigated in ginkgo (Ginkgo biloba L.) leaves from emergence to full size. Under natural conditions, the net photosynthetic rate (P_N), contents of Chl a, Chl b and total soluble proteins and fresh and dry leaf mass gradually increased during leaf expansion. The maximum photochemical efficiency of photosystem (PS) 2 (variable to maximum fluorescence ratio, F_v/F_m) was considerably higher at the early stages of leaf development than in fully expanded leaves. During daily course, only reversible decrease in F_v/F_m was distinguished at various stages, implying that no photo-damage occurred. Absorption flux per cross section (CS) and trapped energy flux per CS were significantly lower in newly expanding leaves compared with fully expanded ones, however, dissipated energy flux per CS was only slightly lower in expanding leaves. The ratio of carotenoids (Car)/Chl decreased gradually during leaf expansion due to increasing Chl content. Moreover, activities of the antioxidant enzymes, such as superoxide dismutase, ascorbate peroxidase, catalase and peroxidase, increased at the early stages of leaf expansion. The appearance of osmiophilic granules in fully expanded leaves further proves that photo-protection is significantly strengthened at the early stages of leaf expansion.     

Ozone-induced changes in photosynthesis and photorespiration of hybrid poplar in relation to the developmental stage of the leaves

Young poplar trees (Populus tremula Michx. x Populus alba L. clone INRA 717-1B4) were subjected to 120 ppb of ozone for 35 days in phytotronic chambers. Treated trees displayed precocious leaf senescence and visible symptoms of injury (dark brown/black upper surface stippling) exclusively observed on fully expanded leaves. In these leaves, ozone reduced parameters related to photochemistry (Chl content and maximum rate of photosynthetic electron transport) and photosynthetic CO(2) fixation [net CO(2) assimilation, Rubisco (ribulose-1,5-bisphosphate carboxylase oxygenase) activity and maximum velocity of Rubisco for carboxylation]. In fully expanded leaves, the rate of photorespiration as estimated from Chl fluorescence was markedly impaired by the ozone treatment together with the activity of photorespiratory enzymes (Rubisco and glycolate oxidase). Immunoblot analysis revealed a decrease in the content of serine hydroxymethyltransferase in treated mature leaves, while the content of the H subunit of the glycine decarboxylase complex was not modified. Leaves in the early period of expansion were exempt from visible symptoms of injury and remained unaffected as regards all measured parameters. Leaves reaching full expansion under ozone exposure showed potential responses of protection (stimulation of mitochondrial respiration and transitory stomatal closure). Our data underline the major role of leaf phenology in ozone sensitivity of photosynthetic processes and reveal a marked ozone-induced inhibition of photorespiration.     

RESPONSE OF LEAF ANATOMY AND PHOTOSYNTHETIC CAPACITY IN ALOCASIA MACRORRHIZA (ARACEAE) TO A TRANSFER FROM LOW TO HIGH LIGHT

Alocasia macrorrhiza plants were grown in 1% and 20% full sunlight, and their leaf anatomical and physiological parameters were measured. Total leaf thickness was 41% greater and mesophyll thickness was 52% greater in high-light leaves than in low-light leaves. This increase in thickness resulted from both increased cell size and number. Maximum leaf photosynthetic capacity was also 66% greater in high- than in low-light leaves. When low-light plants were transferred to high light, the thickness of mature leaves did not increase but the thickness of the first leaf to expand after the transfer was significantly greater than that of the low-light leaves. Thus, only leaves that were still expanding at the time of transfer developed leaf thickness greater than plants remaining in low light. Fully mature leaves showed no change in photosynthetic capacity in response to transfer. Leaves that had just completed expansion at the time of low- to high-light transfer were able to develop slightly higher maximum photosynthetic capacities than older leaves. However, full photosynthetic acclimation to the new light environment did not occur until the second new leaf expanded after transfer. These results are discussed in relation to the timing and mechanisms of whole plant acclimation to increased light.     

Cytokinin-Regulated Mesophyll Cell Division and Expansion during Development of Cucurbita pepo Leaves

The role of cytokinins in the development of mesophyll structure was studied in developing pumpkin Cucurbita pepo L. leaves. Leaves were treated with cytokinins at different stages of growth: when they reached 25 or 50% of their final size (S _max), immediately after leaf growth ceased, and during senescence. At the early stages of leaf development, treatment with exogenous benzyladenine accelerated division of mesophyll cells. At the later stages of development, BA treatment activated expansion of growing cells and those, which have just accomplished their growth. The exogenous cytokinin did not affect the senescent leaf cells. The content of endogenous cytokinins changed during mesophyll development. The juvenile leaves (25% of S _max) were characterized by low level of these phytohormones. In the expanding leaves (50% of S _max), the content of phytohormones increased and decreased when leaf growth ceased. In the senescent leaves, the cytokinin content decreased markedly. It was concluded that the response of mesophyll cells to cytokinin depended on the cell growth phase at the moment of hormone action. Furthermore, in the young leaves, lower cytokinin concentrations were required for division of mesophyll cells in vivo than for cell expansion at the final stage of leaf development.     

Preparation of Escherichia coli 70S ribosomes labeled with 35S

[35S]--70S ribosomes (150 Ci/mmol) were isolated from E. coli MRE-600 cells grown on glucose-mineral media in the presence of [35S] ammonium sulfate. The labeled 30S and 50S subunits were obtained from [35S] ribosomes by centrifugation in a sucrose density gradient of 10--30% under dissociating conditions (0.5 mM Mg2+). The activity of [35S]--70S ribosomes obtained by reassociation of the labeled subunits during poly(U)-dependent diphenylalanine synthesis was not less than 70%. The activity of [35S]--70S ribosomes during poly(U)-directed polyphenylalanine synthesis was nearly the same as that of the standard preparation of unlabeled ribosomes. The 23S, 16S and 5S RNAs isolated from labeled ribosomes as total rRNA contained no detectable amounts of their fragments as revealed by polyacrylamide gel electrophoresis. The [35S] ribosomal proteins isolated from labeled ribosomes were analyzed by two-dimensional gel electrophoresis. The [35S] label was found in all proteins, with the exception of L20, L24 and L33 which did not contain methionine or cysteine residues.     

Avoiding high relative air humidity during critical stages of leaf ontogeny is decisive for stomatal functioning

Plants of several species, if grown at high relative air humidity (RH ≥85%), develop stomata that fail to close fully in case of low leaf water potential. We studied the effect of a reciprocal change in RH, at different stages of leaf expansion of Rosa hybrida grown at moderate (60%) or high (95%) RH, on the stomatal closing ability. This was assessed by measuring the leaf transpiration rate in response to desiccation once the leaves had fully expanded. For leaves that started expanding at high RH but completed their expansion after transfer to moderate RH, the earlier this switch took place the better the stomatal functioning. Leaves initially expanding at moderate RH and transferred to high RH exhibited poor stomatal functioning, even when this transfer occurred very late during leaf expansion. Applying a daily abscisic acid (ABA) solution to the leaves of plants grown at continuous high RH was effective in inducing stomatal closure at low water potential, if done before full leaf expansion (FLE). After FLE, stomatal functioning was no longer affected either by the RH or ABA level. The results indicate that the degree of stomatal adaptation depends on both the timing and duration of exposure to high RH. It is concluded that stomatal functionality is strongly dependent on the humidity at which the leaf completed its expansion. The data also show that the effect of ambient RH and the alleviating role of ABA are restricted to the period of leaf expansion.     

SEASONAL PATTERNS OF LEAF PHOTOSYNTHETIC CAPACITY IN SUCCESSIONAL NORTHERN HARDWOOD TREE SPECIES

Seasonal patterns of leaf photosynthetic capacity and conductance were determined for deciduous hardwood tree species in natural habitats in northern lower Michigan. Leaves of bigtooth aspen and red oak at the top of the canopy had higher maximum CO₂ Exchange Rate (CER) (10–15 μmol m ² s ¹) than leaves of sugar maple, red maple, red oak, and beech growing in the understory (4–5 μmol m ² s ¹). In all leaves, CER measured at light-saturation increased to a maximum near the completion of leaf expansion in early June, was constant until mid-September, and then rapidly declined until leaf death. A similar pattern was seen for CER measured in low light (1.5% full sun). Respiration rate in the dark was highest in young leaves and decreased during leaf expansion; a relatively constant rate was then maintained for the rest of leaf lifespan. The seasonal pattern of the initial slope of the light response of CER paralleled the pattern of light-saturated CER. The initial slope in midsummer ranged from values of 37 to 44 μmol/mol for species in the understory to 51 and 56 μmol/mol for red oak and bigtooth aspen, respectively, at the top of the canopy. Leaf conductance was constant throughout most of leaf lifespan, with some decline occurring in autumn. Leaves at the top of the canopy had higher conductances for water vapor (2–5 mm/s) than leaves in the understory (1–2 mm/s). All species maintained leaf intercellular CO, mole fractions (c,) near 200 uML/L until autumn, when c, increased during leaf senescence.     

Leaf growth and demography of the rheophytic fern Thelypteris angustifolia (Willdenow) Proctor in a Puerto Rican rainforest

Morphological and temporal aspects of the growth of leaves of Thelypteris angustifolia (Willdenow) Proctor are under study along a 400-m section of the Sonadora River at the El Verde Field Station in the Luquillo Experimental Forest of Puerto Rico. Emergence, expansion and growth of leaves of 149 sporophytes have been observed approximately mid-month every January, May and September since September 1991. Results of the first four years of this long-term study of T. angustifolia are reported here. Although they are land plants, sporophytes of T. angustifolia are rheophytic, subject to intermittent flooding and submerged approximately 7% of the time. The pinnate leaves of T. angustifolia exhibit sterile-fertile leaf dimorphy. Only 7.6% of the leaves in the the sample were fertile. The petiole of a fertile leaf was 41% longer than that of a sterile leaf while leaf blades were the same length. Leaves expanded at a mean rate of 1.7 cm per day, maturing in 29.7 days. Mean leaf life span was 10.8 months with a maximum of 26 months. Sterile leaf longevity (11.0 months) was longer than that of the fertile leaf (9.6 months). Leaves were produced at a mean rate of 4.7 leaves per plant per year. Leaf damage occurred on 38% of the leaves observed. Net leaf counts for individual sporophytes ranged from one to eight with a mean of 3.0 leaves per plant. Fertile plants had higher mean leaf counts (4.1) than plants without fertile leaves (2.8). Leaf production, although possibly triggered by increasing daylength, also paralleled seasonal increases in rainfall during the wetter mid-May/September months. Increased leaf mortality occurred during the drier mid-January/May period.     

Powdery mildew of tabacco (Erysiphe cichoracearum DC.)

The natural spread of Erysiphe cichoracearum was assessed weekly on alternate leaves of irrigated and non-irrigated tobacco plants of Kutsaga 51 variety, grown in field plots in 1962-63. Leaf area, air temperature and humidity within the plots, relative turgidity of the leaves and soil moisture were also measured. Leaves emerged over a period of 37 days. A minimum of 29 days elapsed between leaf emergence and infection; irrigation lengthened this period by 2–6 days for leaves 2–6 and shortened it by 2–10 days for leaves 10–18. The duration of the initial resistant phase, in leaves at comparable stalk positions, appeared to be directly proportional to the eventual size of the leaves at reaping. Leaves were not infected until they were almost fully expanded. The longest dry period, when most irrigation water was applied, occurred when most lower leaves (2–8) were fully expanded and already infected; upper leaves (10–18)w ere then still expanding and not yet infected. Irrigation increased infection in all leaves; it increased the growth of the pathogen during dry weather and the subsequent susceptibility of leaves that were still actively expanding but not yet infected. Irrigation increased the percentage of susceptible leaf area infected, of intact plants, threefold and that of topped plants ninefold. Topped plants had less infection than intact ones.     

Survival curves and longevity of the leaves of Alnus japonica var. arguta in Kushiro Marsh

The seasonal changes in leaf emergence and leaf-fall in a Japanese alder stand of the fen in Kushiro Marsh were studied, and survival curves for the leaves were drawn. Leaves collected in litter traps were dried and weighed to study the seasonal changes, peaks in mid-August and late September to October suggested a bimodal annual curve. Study of the seasonal changes in the number of emerged and fallen leaves per shoot revealed a third peak about one month before the August peak, showing a trimodal annual curve. First and second leaves had a longevity of about 40 and 50–60 days, respectively. The longevity increased until the fifth leaf. With the sixth and following leaves, longevity decreased. Leaf size increased with leaf rank, with the first leaf being the smallest. The first leaf had only about 10% and the second leaf only 20% of the area of the fifth leaf. On this basis, the early to mid-July peak in number of fallen leaves was composed of first and second leaves which were smaller and short-lived. The early August and September/October peaks were high in both number and mass of fallen leaves. Compared to reports on Japanese alder of other mountainous districts in Hokkaido, the alder trees of Kushiro Marsh had about the same number of leaves per shoot, but had a season of leaf emergence which was about 6 weeks shorter. In addition, the longevity of the longest-lived fifth leaf was about 30–40 days shorter. The short life span of the leaves could be considered as an adaptive strategy of this species to environmental constraints of its habitat.     

Heteroblasty in the moss,Aphanoregma patens (Physcomitrella patens), results from progressive modulation of a single fundamental leaf developmental programme

Abstract

Leaves at the apex of a mature Aphanoregma patens (Hedw.) Lindb. (Physcomitrella patens (Hedw.) Bruch Schimp. in B.S.G.) gametophore differ markedly in size and form from those at its base. To determine how these differences are produced during development, we first examined qualitative and quantitative differences between successive leaves along the stem and among leaves at different developmental stages. Differences between successive leaves were slight and cumulative. Local changes in cell number and size combined to produce a regularly shaped and approximately bilaterally symmetrical leaf suggesting that cell division and cell expansion are regionally regulated and coordinated at the organ level. The midrib and marginal teeth are discrete characters, which were prefigured by changes in cell shape in leaves that lacked these characters. In leaf primordia, cell proliferation was responsible for most of the changes in leaf form and size early in development and may have continued as cell expansion took over as the primary contributor to leaf growth and morphogenesis. Thus, leaf heteroblasty in Physcomitrella probably results from modulation of a single developmental programme by external and/or internal forces, which alter progressively in intensity as a gametophore grows. We applied exogenous cytokinin and auxin separately to growing cultures to explore their effects on leaf growth. Cytokinin and auxin stimulated leaf cell division and leaf cell elongation, respectively. Also, young upper leaves of gametophores exposed to exogenous auxin closely resembled basal leaves of untreated plants. Therefore, endogenous cytokinins and auxins may be among the modulating internal forces involved in leaf morphogenesis and the establishment of leaf heteroblasty.     

Do small leaves expand faster than large leaves,and do shorter expansion times reduce herbivore damage?

Leaves are most vulnerable to herbivory during expansion. We hypothesised that one factor favouring small leaves could be that smaller-leaved species have shorter expansion times and are therefore exposed to high levels of herbivory for a shorter period than large leaves. In order to test this hypothesis, leaf expansion time and leaf area loss were measured for 51 species from Sydney, Australia. Strong positive correlations were found between leaf length and area and leaf expansion time, confirming that small leaves do expand in a shorter time than large leaves. The amount of leaf area lost was highly variable (from 0.5 to 90% of total leaf area), but was significantly related to both leaf expansion time and log leaf area. The amount of leaf area lost was not significantly correlated with specific leaf area nor with the presence of distasteful substances in the leaves, but was lower on species with hairy expanding leaves.