Leaf life spans in wild perennial herbaceous plants: a survey and attempts at a functional interpretation

Summary Leaf longevity in 29 herbaceous plant species of Central Europe was studied by inspecting tagged leaves at weekly intervals. About half of the species are elements of the lowland meadow flora, the other half comprises a representative sample of species from the highest sites where vascular plants grow in the Alps. Shaded and water-stressed sites were avoided. Overall mean leaf longevity did not differ significantly between sites and amounted to 71±5 days at low and 68±4 days at high altitude. Leaf life spans ranged (with no clear altitudinal trend) from 41 to 95 days. Low-altitude forbs and grasses produced several leaf cohorts during their growth period, while most alpine species produced only one. Correlations were found between leaf duration and percent nitrogen content and carbon-cost/carbon-gain ratios, but not with leaf dry mass per unit leaf area and photosynthetic capacity alone. As leaf life spans increase, more C tends to be invested per unit CO₂ uptake and less N is invested per unit invested C. Thus, mass relationships rather than area relationships seem to be linked to leaf life span in these species, suggesting that leaf duration is associated with properties other than the efficiency of light utilization (e.g. mechanical strength, herbivory or pathogen resistance). It seems that the explanations of leaf duration that have been developed for evergreen/deciduous plants and for plants along steep light gradients do not apply to the variable life spans in leaves of perennial herbaceous plants of open habitats.     

Ecological significance of leaf life span among Central European grass species

Interspecific variation in leaf life span reflects the variation in nutrient conservation ability among different plant species and is considered to be associated with nutrient availability in the characteristic habitat. As defoliation interferes with nutrient conservation by the long-lived leaves, we hypothesized that disturbance rate is another important environmental factor working as a selective force on interspecific variation in leaf life span. In order to investigate this, we measured leaf life span of 32 grass species in mature garden-grown individuals. Variation in leaf life span was compared to measured leaf traits, to available data on species occurrence along gradients of nutrient availability and disturbance, and to published relative growth rates of the species. Leaf life span was associated positively with leaf tissue mass density and negatively with specific leaf area. Leaf life span correlated negatively with the disturbance rate in the characteristic habitat of a species, but not with nutrient availability. The latter relationship did not come about due to the long leaf life spans of species from nutrient-rich habitats with a relatively low disturbance rate, and to some extent also due to the short leaf life spans of annual species from relatively nutrient-poor sites. We conclude that although leaf longevity is an important means of reducing nutrient losses, this is a selective advantage only if the plant is not subjected to frequent defoliation. The frequently postulated association between leaf life span of a species and nutrient availability in its characteristic habitat may occur among species of habitats with positively correlated nutrient availability and disturbance rate. Leaf life span is negatively associated with seedling RGR, but there may be deviations in this relationship due to species with contrasting characteristics at seedling stage and at maturity.     

A cost-benefit analysis of acclimation to low irradiance in tropical rainforest tree seedlings: leaf life span and payback time for leaf deployment

The maintenance in the long run of a positive carbon balance under very low irradiance is a prerequisite for survival of tree seedlings below the canopy or in small gaps in a tropical rainforest. To provide a quantitative basis for this assumption, experiments were carried out to determine whether construction cost (CC) and payback time for leaves and support structures, as well as leaf life span (i) differ among species and (ii) display an irradiance-elicited plasticity. Experiments were also conducted to determine whether leaf life span correlates to CC and payback time and is close to the optimal longevity derived from an optimization model. Saplings from 13 tropical tree species were grown under three levels of irradiance. Specific-CC was computed, as well as CC scaled to leaf area at the metamer level. Photosynthesis was recorded over the leaf life span. Payback time was derived from CC and a simple photosynthesis model. Specific-CC displayed only little interspecific variability and irradiance-elicited plasticity, in contrast to CC scaled to leaf area. Leaf life span ranged from 4 months to >26 months among species, and was longest in seedlings grown under lowest irradiance. It was always much longer than payback time, even under the lowest irradiance. Leaves were shed when their photosynthesis had reached very low values, in contrast to what was predicted by an optimality model. The species ranking for the different traits was stable across irradiance treatments. The two pioneer species always displayed the smallest CC, leaf life span, and payback time. All species displayed a similar large irradiance-elicited plasticity.     

Geography of Pinus elliottii Engelm. and Pinus palustris Mill. leaf life-spans in the southeastern U.S.A.

Aim Past studies have investigated differences in leaf life‐spans between deciduous and evergreen species. Environmental controls such as light, temperature, and nutrient and moisture availability explain differences in leaf life‐spans between species. This study examined intraspecific leaf life‐spans across climate and nutrient gradients within the geographical range of Pinus palustris Engelm (longleaf pine) and Pinus elliottii Mill. (slash pine). Location Five study areas in the southeastern United States were selected along the north–south geographical range of Pinus elliottii and Pinus palustris. Methods Leaf life‐span was calculated based on stand inventories and annual litterfall totals for each site, and allometric relationships between d.b.h. and foliar biomass. Results Leaf life‐span of P. elliottii ranged from 1.28 to 1.95 years between sites. Leaf life‐span of P.palustris varied by nearly a factor of 5 between the study site with the lowest and highest value (0.58–2.49 years). anova indicated that leaf life‐spans of P. elliottii were not significantly different among sites. In contrast, anova indicated a significant difference for P. palustris leaf life‐spans among sites (P < 0.05). The Tukey multiple comparisons tests showed that 2 study areas were the only pair of P. palustris sites with a significant difference in leaf life‐spans. Main conclusions The geographical variation in leaf life‐spans between two species illustrates the different phenotypic responses to environmental controls. The variation in leaf life‐spans by individuals of P. palustris across a geographical range illustrated in this study suggests that P. palustris may exhibit a greater phenotypic plasticity than P. elliottii.     

Leaf demography of <Emphasis Type="Italic">Festuca pallens</Emphasis> in dry grassland communities

Leaf blade parameters and leaf demography of Festuca pallens Host were studied in two types of dry grasslands. The field work was carried out in the Považsky Inovec Mts (Western Carpathians) during 1993–1995. The permanent plot in the Poo badensis-Festucetum pallentis was located on a steep, strongly eroded S-facing slope covered with dolomite outcrops, scree and sparse vegetation (20%) dominated by Festuca pallens. The permanent plot in the Festuco pallentis-Caricetum humilis was located on the even ridge plateau with shallow stony soil and vegetation covering about 70% dominated by Carex humilis and Festuca pallens. In comparison to other grasses Festuca pallens had a very low rate of leaf turnover. The highest leaf birth rates and the lowest leaf death rates were observed in June. Leaf mortality was uniformly distributed in time without a distinct minimum or maximum. For the surviving tillers the leaf production exceeded the leaf mortality during the whole growing season. The steady net gain of leaves in tillers was not interrupted by the parallel process of tillering. Among the leaf cohorts the leaves produced in May had the longest leaf blades. Leaves grew during the whole year. The winter cold and summer drought might slow down the growth rate or interrupt the growth. The growth of a leaf blade took five to eight weeks. Leaf life span was estimated to 150–200 days (time from leaf appearance at the apex to the complete loss of its green assimilating parts). In comparison to other grasses Festuca pallens belongs to the species with the longest leaf life span. The effect of environmental factors on leaf demography was followed by the comparison of two populations belonging to two phytosociological associations differing mostly in habitat xericity. Differences were revealed in the following characteristics: length of leaf blade in cohorts born during May and June, maximum length of a leaf blade in a tiller and daily increments in May and June. The course of leaf natality and mortality was similar in the studied populations.     

A Cost-Benefit Analysis of Leaves of Eight Australian Savanna Tree Species of Differing Leaf Life-Span

Cost-benefit analysis of foliar construction and maintenance costs and of carbon assimilation of leaves of differing life-span were conducted using two evergreen, three semi-deciduous, and three deciduous tree species of savannas of north Australia. Rates of radiant-energy-saturated CO₂ assimilation (P _max) and dark respiration were measured and leaves were analysed for total nitrogen, fat, and ash concentrations, and for heat of combustion. Specific leaf area, and leaf N and ash contents were significantly lower in longer-lived leaves (evergreen) than shorter-lived leaves (deciduous) species. Leaves of evergreen species also had significantly higher heat of combustion and lower crude fat content than leaves of deciduous species. On a leaf area basis, P _max was highest in leaves of evergreen species, but on a leaf dry mass basis it was highest in leaves of deciduous species. P _max and total Kieldahl N content were linearly correlated across all eight species, and foliar N content was higher in leaves of deciduous than evergreen species. Leaf construction cost was significantly higher and maintenance costs were lower for leaves of evergreen than deciduous species. Maintenance and construction costs were linearly related to each other across all species. Leaves of evergreen species had a higher cost-benefit ratio compared to leaves of deciduous species but with longer lived leaves, the payback interval was longer in evergreen than deciduous species. These results support the hypotheses that longer lived leaves are more expensive to construct than short-lived leaves, and that a higher investment of N into short-lived leaves occurs which supports a higher P _max over a shorter payback interval.     

Response to grazing after nine years of cattle exlusion in a Flooding Pampa grassland,Argentina

This paper reports on changes induced by the introduction of cattle in a grassland that had remained ungrazed for 9 yr, in comparison with two adjacent grasslands: one that remained enclosed and one that has been continuously subject to grazing. Basal cover was measured on 25 interception lines, each 1 m long, three times during one year. The variables studied were: total cover, cover of grasses and dicots, cover of creeping grasses, floristic composition, and dissimilarity among sites. At the first sampling, 2 yr after cattle re-introduction, the newly grazed site was more similar to the ungrazed than to the grazed site. The newly grazed site had very low cover of dicots; the species of dicots present were different from those found in the continuously grazed area. Creeping grasses had higher cover in the newly grazed site than in the other sites, and continued to increase. At the last sampling, one year later, the newly grazed site had become more similar to the contiuously grazed site. Only after 5 yr of cattle grazing the exotic dicots that were dominant in the continuously grazed site, were recorded in the re-opened site. The absence of propagules of these species or the absence of safe sites may account for this delayed invasion.     

Red reveals branch die-back in Norway maple Acer platanoides

Background and Aims

Physiological data suggest that autumn leaf colours of deciduous trees are adaptations to environmental stress. Recently, the evolution of autumn colouration has been linked to tree condition and defence. Most current hypotheses presume that autumn colours vary between tree individuals. This study was designed to test if within-tree variation should be taken into account in experimental and theoretical research on autumn colouration.

Methods

Distribution of red autumn leaf colours was compared between partially dead and vigorous specimens of Norway maple (Acer platanoides) in a 3-year study. In August, the amount of reddish foliage was estimated in pairs of partially dead and control trees. Within-tree variation in the distribution of reddish leaves was evaluated. Leaf nitrogen and carbon concentrations were analysed.

Key Results

Reddish leaf colours were more frequent in partially dead trees than in control trees. Reddish leaves were evenly distributed in control trees, while patchiness of red leaf pigments was pronounced in partially dead trees. Large patches of red leaves were found beneath or next to dead tree parts. These patches reoccurred every year. Leaf nitrogen concentration was lower in reddish than in green leaves but the phenomenon seemed similar in both partially dead and control trees.

Conclusions

The results suggest that red leaf colouration and branch condition are interrelated in Norway maple. Early reddish colours may be used as an indication of leaf nitrogen and carbon levels but not as an indication of tree condition. Studies that concentrate on entire trees may not operate at an optimal level to detect the evolutionary mechanisms behind autumnal leaf colour variation.Key words: Acer platanoides, Norway maple, branch die-back, coevolution hypothesis, leaf senescence, patchy distribution, red leaf pigments, tree condition, within-tree variation     

Longevity, lignin content and construction cost of the assimilatory organs of Nepenthes species

Background and Aims

This study examined level of causal relationships amongst functional traits in leaves and conjoint pitcher cups of the carnivorous Nepenthes species.

Methods

Physico-chemical properties, especially lignin content, construction costs, and longevity of the assimilatory organs (leaf and pitcher) of a guild of lowland Nepenthes species inhabiting heath and/or peat swamp forests of Brunei, northern Borneo were determined.

Key Results

Longevity of these assimilatory organs was linked significantly to construction cost, lignin content and structural trait of tissue density, but these effects are non-additive. Nitrogen and phosphorus contents (indicators of Rubisco and other photosynthetic proteins), were poor predictors of organ longevity and construction cost, suggesting that a substantial allocation of biomass of the assimilatory organs in Nepenethes is to structural material optimized for prey capture, rigidity and escape from biotic and abiotic stresses rather than to light interception. Leaf payback time – a measure of net carbon revenue – was estimated to be 48–60 d. This is in line with the onset of substantial mortality by 2–3 months of tagged leaves in many of the Nepenthes species examined. However, this is a high ratio (i.e. a longer minimum payback time) compared with what is known for terrestrial, non-carnivorous plants in general (5–30 d).

Conclusions

It is concluded that the leaf trait bivariate relationships within the Nepenthes genus, as in other carnivorous species (e.g. Sarraceniaceae), is substantially different from the global relationship documented in the Global Plant Trait Network.Key words: Botanical carnivory, carbon gain, functional traits, leaf chemistry, leaf lifespan, leaf mass per unit area, Nepenthes, pitcher, payback time     

Monocot leaves are eaten less than dicot leaves in tropical lowland rain forests: correlations with toughness and leaf presentation

Background and Aims

In tropical lowland rain forest (TLRF) the leaves of most monocots differ from those of most dicots in two ways that may reduce attack by herbivores. Firstly, they are tougher. Secondly, the immature leaves are tightly folded or rolled until 50–100 % of their final length. It was hypothesized that (a) losses of leaf area to herbivorous invertebrates are generally greatest during leaf expansion and smaller for monocots than for dicots, and (b) where losses after expansion are appreciable any difference between monocots and dicots then is smaller than that found during expansion.

Methods

At six sites on four continents, estimates were made of lamina area loss from the four most recently mature leaves of focal monocots and of the nearest dicot shoot. Measurements of leaf mass per unit area, and the concentrations of water and nitrogen were made for many of the species. In Panama, the losses from monocots (palms) and dicots were also measured after placing fully expanded palm leaflets and whole dicot leaves on trails of leaf-cutter ants.

Key Results

At five of six sites monocots experienced significantly smaller leaf area loss than dicots. The results were not explicable in terms of leaf mass per unit area, or concentrations of water or nitrogen. At only one site was the increase in loss from first to fourth mature leaf significant (also large and the same in monocots and dicots), but the losses sustained during expansion were much smaller in the monocots. In the leaf-cutter ant experiment, losses were much smaller for palms than for dicots.

Conclusions

The relationship between toughness and herbivory is complex; despite the negative findings of some recent authors for dicots we hypothesize that either greater toughness or late folding can protect monocot leaves against herbivorous insects in tropical lowland rain forest, and that the relative importance varies widely with species. The difficulties of establishing unequivocally the roles of leaf toughness and leaf folding or rolling in a given case are discussed.Key words: anti-herbivore defences, dicots, herbivory, leaf folding, leaf rolling, leaf toughness, monocots, palms, tropical rain forest     

Leaf dynamics of seedlings of rain forest species in relation to canopy gaps

Summary Leaf dynamics of eight tropical rain forest species seedlings was studied in three environments: the shaded forest understorey, a small gap of ±50 m², and a large gap of ±500 m². Leaf production rate and leaf loss rate were enhanced in gaps, and a large gap resulted in larger increases than a small gap. For most species net leaf gain rate was larger in gaps, although this rate was not always largest in the large gap. Leaf loss decreased, and leaf survival percentages increased with increasing shade tolerance of species, indicating a slower leaf turnover for more shade tolerant species. Leaf area growth rate was only partly determined by net leaf gain rate. Ontogenetic effects on leaf size were also important, especially in the large gap. Species which possessed leaves with high specific leaf weight (SLW) showed lower leaf loss rates and higher leaf survival percentages than species with low SLW leaves. Leaf life span seemed to be related to leafcost per unit area. The relation of specific patterns in leaf production and leaf loss to the regeneration mode of the species is briefly discussed.     

Experimental Evaluation of Herbivory on Live Plant Seedlings by the Earthworm Lumbricus terrestris L. in the Presence and Absence of Soil Surface Litter

Background

Recent studies suggested that the earthworm Lumbricus terrestris might act as a seedling predator by ingesting emerging seedlings, and individuals were observed damaging fresh leaves of various plant species in the field. To evaluate the significance of herbivore behavior of L. terrestris for plant and earthworm performance we exposed 23- to 33-days-old seedlings of six plant species to earthworms in two microcosm experiments. Plants belonged to the three functional groups grasses, non-leguminous herbs, and legumes. Leaf damage, leaf mortality, the number of leaves as well as mortality and growth of seedlings were followed over a period of up to 26 days. In a subset of replicates 0.1 g of soil surface litter of each of the six plant species was provided and consumption was estimated regularly to determine potential feeding preferences of earthworms.

Results

There was no difference in seedling growth, the number of live seedlings and dead leaves between treatments with or without worms. Fresh leaves were damaged eight times during the experiment, most likely by L. terrestris, with two direct observations of earthworms tearing off leaf parts. Another nine leaves were partly pulled into earthworm burrows. Lumbricus terrestris preferred to consume legume litter over litter of the other plant functional groups. Earthworms that consumed litter lost less weight than individuals that were provided with soil and live plants only, indicating that live plants are not a suitable substitute for litter in earthworm nutrition.

Conclusion

Our results demonstrate that L. terrestris damages live plants; however, this behavior occurs only rarely. Pulling live plants into earthworm burrows might induce microbial decomposition of leaves to make them suitable for later consumption. Herbivory on plants beyond the initial seedling stage may only play a minor role in earthworm nutrition and has limited potential to influence plant growth.     

Leaf traits and leaf life spans of two xeric-adapted palmettos

Plants of nutrient-poor, arid environments often have leaf traits that include small size, sclerophylly, long life span, low nutrient concentration, and low photosynthetic rate. Hence, the success of two large-leaved palmettos in peninsular Florida's seasonally xeric, nutrient-impoverished uplands seems anomalous, given that their leaves are orders of magnitude larger than the leaves of sympatric species. An examination of a 16-yr data set of leaf traits and leaf life spans across four vegetative associations differing in available light showed that Serenoa repens and Sabal etonia had low rates of leaf production coupled with long leaf life spans reaching 3.5 yr in heavily shaded plants. The adaptation of these palmettos to xeric, nutrient-poor habitats has generated dwarf statures, diminished leaf sizes and numbers, increased leaf life spans, and reduced rates of leaf production relative to other palms and congeners of more mesic sites. Leaf and petiole size, plant leaf canopy area, and leaf life span increased in both palmettos with decreasing available light, helping to compensate for reduced photosynthetic rates under shaded conditions and for the high leaf construction costs of the large, thick palmetto leaves. Large leaf size in these palmettos, likely due to phylogenetic conservatism, is compensated by other leaf traits (e.g., heavily cutinized epidermises, thick laminas) that increase survival in seasonally xeric, nutrient-impoverished environments.     

Matter-economical roles of the evergreen foliage ofAucuba japonica,an understory shrub in the warm-temperate region of Japan

Leaf demography and productivity ofAucuba japonica, an understory shrub in the warm-temperate region, were examined and dry matter economy was analyzed to evaluate the roles of the evergreen foliage. Turnover of leaves occurred during a short period in spring. The mean leaf life span was about 2.6 years. Annual NAR (net assimilation rate) of each sample shoot was calculated from the biomass and the total dead mass estimated from scars of leaves and floral parts. The average NAR was 1.34±0.22 g·g⁻¹·yr⁻¹. The ratio of dry matter produced by leaves during their whole life span to the initial investment was 3.45±0.37. The annual NAR calculated for individual plants was negatively related to the life span of their leaves. The seasonal change in SLW (specific leaf weight) showed that the reserve material in leaves was accumulated from autumn to early spring and was consumed for the growth of new organs in the following season. The dry matter withdrawn in spring from the overwintering foliage amounted to 40% of dry mass of the new organs developed.     

Plant diversity patterns in semi-natural grasslands along an elevational gradient in southern Norway

Leaf demography and growth of six common, co-occurring woody plant species that varied in stature (tree vs. shrub) and leaf texture (sclerophyllous, coriaceous, malacophyllous) were examined in a subtropical savanna parkland in southern Texas, USA. We tested the hypotheses that, (a) leaves of plants with evergreen canopies would have longer life spans than those of deciduous species; (b) supplementation of soil moisture would decrease leaf life span in both evergreen and deciduous species; (c) species responses to increased soil moisture availability would be inversely related to leaf longevity; and (d) deciduous growth forms would exhibit a greater growth response to increased soil moisture availability than their evergreen counterparts.A variety of seasonal leaf habits (evergreen, winter-deciduous and summer-deciduous canopies) and leaf life spans (median = 66 to 283 days) were represented by the targeted species, but there was no clear relationship between seasonal leaf habit and leaf longevity. Among species with evergreen canopies, median leaf longevity ranged from short (Zanthoxylum fagara = 116 days; Condalia hookeri = 158 days) to long (Berberis trifoliolata = 283 days) but did not exceed 1 yr. In fact, leaf longevity in evergreen shrubs was often comparable to, or shorter than, that of species with deciduous canopies (Ziziphus obtusifolia = 66 days; Diospyros texana = 119 days; Prosopis glandulosa = 207 days). Augmentation of surface soil moisture had no detectable effect on median leaf life span in any species and there was no clear relationship between leaf longevity and species growth responses to irrigation. Contrary to expectations, species with evergreen canopies responded to irrigation by producing more leaf biomass, longer shoots and more leaf cohorts/year than did deciduous species.Species differences in the annual cycle of leaf initiation, leaf longevity and canopy development, combined with contrasts in root distributions and a highly variable climate, may allow for spatial and temporal partitioning of resources and hence, woody species coexistence and diversity in this system. However, the lack of expected relationships between leaf longevity, leaf habit and plant responses to resource enhancement suggests that structure-function relationships and functional groupings developed in strongly seasonal environments cannot be applied with confidence to these subtropical savannas and thorn woodlands.     

Responses of Tropical Understory Plants to a Severe Drought: Tolerance and Avoidance of Water Stress1

Shade-tolerant understory shrubs and subcanopy trees constitute most of the woody species in Neotropical moist forest, but studies demonstrating physiological differences among these species are few. Shade-tolerant species that coexist in the forest understory exhibit differences in leaf life span that have been associated with variation in physiological traits. We hypothesized that water relations of understory species with widely divergent leaf life spans differ in response to drought. Although severe drought is infrequent in Neotropical moist forest, we studied the water relations of shade-tolerant understory species with short or long leaf life spans during the severe 1991-1992 dry season on Barro Colorado Island, Panama. The predawn leaf water potential declined to -2.8 and -3.6 MPa during the dry season in Hybanthus prunifolius and Psychotria horizontalis, respectively, two species with short leaf life spans, but remained above -1.3 MPa in two species with long leaf life spans, Swartzia simplex and Ouratea lucens. The midday leaf water potential dropped as low as -3.4 and -4.5 MPa for H. prunifolius and P. horizontalis, respectively. The osmotic potential of H. prunifolius and P. horizontalis and another species with short leaf life span, Alms blackiana, decreased early in the dry season, a period during which all three had substantially negative predawn water potential. In contrast, the osmotic potential of S. Simplex, O. lucens, and Licania platypus, a third species with long leaf life span, declined late in the dry season, even though we observed little change in predawn water potential for S. simplex and O. lucens. We conclude that the variable and potentially severe dry season in Neotropical moist forest can be sufficiently intense to severely limit soil moisture availability for understory plants. H. prunifolius and P. horizontalis tolerated dehydration, whereas S. simplex and O. lucens postponed dehydration.     

Leaf physiology does not predict leaf habit; examples from tropical dry forest

Leaf structure and physiology are thought to be closely linked to leaf longevity and leaf habit. Here we compare the seasonal variation in leaf hydraulic conductance (k_leaf) and water potential of two evergreen tree species with contrasting leaf life spans, and two species with similar leaf longevity but contrasting leaf habit, one being deciduous and the other evergreen. One of the evergreen species, Simarouba glauca, produced relatively short-lived leaves that maintained high hydraulic conductance year round by periodic flushing. The other evergreen species, Quercus oleoides, produced longer-lived leaves with lower k_leaf and as a result minimum leaf water potential was much lower than in S. glauca (–2.8 MPa vs –1.6 MPa). Associated with exposure to lower water potentials, Q. oleoides leaves were harder, had a higher modulus of elasticity, and were less vulnerable to cavitation than S. glauca leaves. Both species operate at water potentials capable of inducing 20 (S. glauca) to 50% (Q. oleoides) loss of k_leaf during the dry season although no evidence of cumulative losses in k_leaf were observed in either species suggesting regular repair of embolisms. Leaf longevity in the deciduous species Rhedera trinervis is similar to that of S. glauca, although maximum k_leaf was lower. Furthermore, a decline in leaf water potential at the onset of the dry season led to cumulative losses in k_leaf in R. trinervis that culminated in leaf shedding.     

Periodicity of leaf growth and leaf dry mass changes in the evergreen and deciduous species of Southern Assam,India

The study described patterns of leaf dry mass change, leaf mass per area (LMA), relative growth rate and leaf life span (LL) for 14 evergreen and 7 deciduous species of a tropical forest of Southern Assam, India. Leaf expansion in both the groups was, in general, completed before June (i.e. well before the onset of monsoon rains). Although leaf dry mass during leaf initiation phase was significantly higher (P < 0.01) in evergreen species than in deciduous species, at the time of full leaf expansion, average leaf dry mass relative to the peak leaf dry mass, realised by the evergreen species was lower (66 %) than for deciduous species (76 %). Leaf dry mass increase in both groups continued after leaf full expansion. Evergreen species had a longer leaf dry mass steady phase than deciduous species (2–6 vs 2–3 months). Average LMA of mature leaves for evergreen species (77.43 g m^?2) was significantly greater than that of deciduous species (48.43 g m^?2). LL ranged from 165 days in Gmelina arborea (deciduous) to 509 days in Dipterocarpus turbinatus (evergreen). LMA was correlated positively with LL, indicating that evergreen species with higher leaf construction cost retain leaves for longer period to pay back. The average leaf dry mass loss before leaf shedding was greater (P < 0.01) for deciduous species (30.29 %) than for evergreen species (18.31 %). Although the cost of leaf construction in deciduous species was lower than for evergreen species, they replace leaves at a faster rate. Deciduous species perhaps compensate the cost involved in faster leaf replacement through higher reabsorption of dry mass during senescence, which they remobilise to initiate growth in the following spring when soil resources remain limiting.     

Leaf life span of floating-leaved plants

Photosynthetic capacity of floating-leaved plants is relatively high comparable with terrestrial herbaceous plants, though floating-leaved plants have a much smaller biomass with a leaf area index seldom exceeding 2m²m^-2. Their rather small biomass accumulation is related to higher turnover of leaf biomass or shorter leaf life span. Life span of floating leaves reported in the literature ranged mostly from 13 to 35 days, shorter than that of any other groups of herbaceous macrophytes. Floating-leaved plants are known to show considerably high plasticity in their leaf form. Leaf life span could be prolonged for Nymphoides peltata (Gmel.) O. Kuntze grown in a terrestrial environment and for emergent leaves of Nelumbo nucifera Gaertn. Their short leaf life span seems to be closely related to the fact that old leaves covered by newly formed ones are inevitably compelled to be submerged and lose their function as a photosynthetic apparatus.Abbreviations LAI leaf area index - PFD photosynthetic photon flux density     

Light and growth temperature alter carbon isotope discrimination and estimated bundle sheath leakiness in C<Subscript>4</Subscript> grasses and dicots

We combined measurements of short-term (during gas exchange) and long-term (from plant dry matter) carbon isotope discrimination to estimate CO₂ leakiness from bundle sheath cells in six C₄ species (three grasses and three dicots) as a function of leaf insertion level, growth temperature and short-term irradiance. The two methods for determining leakiness yielded similar results (P > 0.05) for all species except Setaria macrostachya, which may be explained by the leaf of this species not being accommodating to gas exchange. Leaf insertion level had no effect on leakiness. At the highest growth temperature (36°C) leakiness was lower than at the two lower growth temperatures (16°C and 26°C), between which no differences in leakiness were apparent. Higher irradiance decreased leakiness in three species, while it had no significant effect on the others (there was an opposite trend in two species). The inverse response to increasing irradiance was most marked in the two NAD-ME dicots (both Amaranthus species), which both showed almost 50% leakiness at low light (300 μmol quanta m⁻² s⁻¹) compared to about 30% at high light (1,600 μmol quanta m⁻² s⁻¹). NADP-ME subtype grasses had lower leakiness than NAD-ME dicots. Although there were exceptions, particularly in the effect of irradiance on leakiness in Sorghum and Boerhavia, we conclude that conditions favourable to C₄ photosynthesis (high temperature and high light) lead to a reduction in leakiness.