Overwintering leaves of a forest-floor fern, Dryopteris crassirhizoma (Dryopteridaceae): a small contribution to the resource storage and photosynthetic carbon gain

BACKGROUND AND AIMS: Dryopteris crassirhizoma is a semi-evergreen fern growing on the floor of deciduous forests. The present study aimed to clarify the photosynthetic and storage functions of overwintering leaves in this species. METHODS: A 2-year experiment with defoliation and shading of overwintering leaves was conducted. Photosynthetic light response was measured in early spring (for overwintering leaves) and summer (for current-year leaves). KEY RESULTS: No nitrogen limitation of growth was detected in plants subjected to defoliation. The number of leaves, their size, reproductive activity (production of sori) and total leaf mass were not affected by the treatment. The defoliation of overwintering leaves significantly reduced the bulk density of rhizomes and the root weight. The carbohydrates consumed by the rhizomes were assumed to be translocated for leaf production. Photosynthetic products of overwintering leaves were estimated to be small. CONCLUSION: Overwintering leaves served very little as nutrient-storage and photosynthetic organs. They partly functioned as a carbon-storage organ but by contrast to previous studies, their physiological contribution to growth was found to be modest, probably because this species has a large rhizome system. The small contribution of overwintering leaves during the short-term period of this study may be explained by the significant storage ability of rhizomes in this long-living species. Other ecological functions of overwintering leaves, such as suppression of neighbouring plants in spring, are suggested.     

Carbon and nitrogen partitioning in the biennial monocarp Arctium tomentosum Mill

Summary Growth and nitrogen partitioning were investigated in the biennial monocarp Arctium tomentosum in the field, in plants growing at natural light conditions, in plants in which approximately half the leaf area was removed and in plants growing under 20% of incident irradiation. Growth quantities were derived from splined cubic polynomial exponential functions fitted to dry matter, leaf area and nitrogen data.Main emphasis was made to understanding of the significance of carbohydrate and nitrogen storage of a large tuber during a 2-years' life cycle, especially the effect of storage on biomass and seed yield in the second season. Biomass partitioning favours growth of leaves in the first year rosette stage. Roots store carbohydrates at a constant rate and increase storage of carbohydrates and nitrogen when the leaves decay at the end of the first season. In the second season the reallocation of carbohydrates from storage is relatively small, but reallocation of nitrogen is very large. Carbohydrate storage just primes the growth of the first leaves in the early growing season, nitrogen storage contributes 20% to the total nitrogen requirement during the 2nd season. The efficiency of carbohydrate storage for conversion into new biomass is about 40%. Nitrogen is reallocated 3 times in the second year, namely from the tuber to rosette leaves and further to flower stem leaves and eventually into seeds. The harvest index for nitrogen is 0.73, whereas for biomass it is only 0.19.     

Temporal variation in leaf nitrogen partitioning of a broad-leaved evergreen tree, <Emphasis Type="Italic">Quercus myrsinaefolia</Emphasis>

We examined temporal changes in the amount of nitrogenous compounds in leaves from the outer and inner parts of the crown of Quercus myrsinaefolia growing in a seasonal climate. Throughout the leaf life span, metabolic protein and Rubisco content closely correlated with total nitrogen content, while structural protein content was relatively stable after full leaf expansion. Chlorophyll content was affected by shading as well as total nitrogen content in outer leaves that were overtopped by new shoots in the second year. Outer leaves showed a large seasonal variation in photosynthetic nitrogen-use efficiency (PNUE; the light-saturated photosynthetic rate per unit leaf nitrogen content) during the first year of their life, with PNUE decreasing from the peak in summer towards winter. Outer and inner leaves both showed age-related decline in PNUE in the second year. There were no such drastic changes in leaf nitrogen partitioning that could explain seasonal and yearly variations in PNUE. Nitrogen resorption occurred in overwintering leaves in spring. Metabolic protein explained the majority of nitrogen being resorbed, whereas structural protein, which was low in degradability, contributed little to nitrogen resorption.     

Deciduous and evergreen trees differ in juvenile biomass allometries because of differences in allocation to root storage

Background and Aims

Biomass partitioning for resource conservation might affect plant allometry, accounting for a substantial amount of unexplained variation in existing plant allometry models. One means of resource conservation is through direct allocation to storage in particular organs. In this study, storage allocation and biomass allometry of deciduous and evergreen tree species from seasonal environments were considered. It was expected that deciduous species would have greater allocation to storage in roots to support leaf regrowth in subsequent growing seasons, and consequently have lower scaling exponents for leaf to root and stem to root partitioning, than evergreen species. It was further expected that changes to root carbohydrate storage and biomass allometry under different soil nutrient supply conditions would be greater for deciduous species than for evergreen species.

Methods

Root carbohydrate storage and organ biomass allometries were compared for juveniles of 20 savanna tree species of different leaf habit (nine evergreen, 11 deciduous) grown in two nutrient treatments for periods of 5 and 20 weeks (total dry mass of individual plants ranged from 0·003 to 258·724 g).

Key Results

Deciduous species had greater root non-structural carbohydrate than evergreen species, and lower scaling exponents for leaf to root and stem to root partitioning than evergreen species. Across species, leaf to stem scaling was positively related, and stem to root scaling was negatively related to root carbohydrate concentration. Under lower nutrient supply, trees displayed increased partitioning to non-structural carbohydrate, and to roots and leaves over stems with increasing plant size, but this change did not differ between leaf habits.

Conclusions

Substantial unexplained variation in biomass allometry of woody species may be related to selection for resource conservation against environmental stresses, such as resource seasonality. Further differences in plant allometry could arise due to selection for different types of biomass allocation in response to different environmental stressors (e.g. fire vs. herbivory).     

Non-structural carbohydrates and nitrogen dynamics in mediterranean sub-shrubs: an analysis of the functional role of overwintering leaves

Previous studies have led to contrasting results about the role of overwintering leaves as storage sites, which is related to leaf longevity and life-form. The aim of this study was to evaluate the functional role of the leaves of four species of Mediterranean sub-shrubs, with different leaf phenology, as sources of nitrogen (N) and non-structural carbohydrates (NSC) for shoot growth. The seasonal dynamics of the concentrations and pools of N and NSC were assessed monthly in the leaves and woody organs of each species. Overwintering and spring leaves served as N and NSC sources for shoot growth in the evergreen species analyzed, providing up to 73 % and 324 % of the N demand for spring and autumn growth, respectively. Excess autumn N was stored in woody structures which contributed to the N and NSC requirements of spring growth. In the winter deciduous species, woody organs were the main N source for spring growth, while current photosynthesis from immature brachyblasts seemed to be the main carbon (C) source. Due to their short lifespan, overwintering and spring leaves did not show several translocation processes throughout their life time, their contribution to new growth being made during senescence. The successive exchange of leaf cohorts displayed by Mediterranean sub-shrubs might serve as a mechanism to recycle N and C between consecutive cohorts as plants perform the pheno-morphological changes needed to adapt their morphology to the seasonality of their environment.     

Uptake,demand and internal cycling of nitrogen in saplings of Mediterranean<Emphasis Type="Italic"> Quercus</Emphasis> species

Nitrogen uptake, nitrogen demand and internal nitrogen cycling were studied to address the question of the importance of nutrient storage in Quercus species with contrasting leaf longevities. We carried out this study at the whole-plant level with young trees (2-4 years old) of three Mediterranean Quercus species: the evergreen Q. ilex, the marcescent/evergreen Q. faginea, and the deciduous Q. pyrenaica. Seasonal dynamics of nitrogen in all compartments of the plant were followed for 3 years. Nitrogen losses were measured through litter production, herbivory and fine root shedding. Nitrogen uptake was estimated using increments of nitrogen plant content plus accumulative nitrogen losses. Nitrogen uptake was limited to a few months during late winter and spring. Before budbreak, acquired nitrogen was stored in old-leaf cohorts of evergreen and woody compartments. After budbreak, Quercus species relied first on soil uptake and second on nitrogen retranslocation to supply new growth requirements. However, in most cases we found a high asynchrony between nitrogen demand by growing tissues and soil supply, which determined a strong nitrogen retranslocation up to 88.4% of the nitrogen demand throughout leaf expansion. Except for the first year after planting, the above- and underground woody fractions provided more nitrogen to the new tissues than the old leaf cohorts. Differences in the benefit of nitrogen withdrawn from senescent and old leaves were not found between species. We conclude that sink/source interaction strength was determined by differences between nitrogen demand and uptake, regulating internal nutrient cycling at the whole plant level.     

Differences in response to simulated herbivory between <Emphasis Type="Italic">Quercus crispula</Emphasis> and <Emphasis Type="Italic">Quercus dentata</Emphasis>

To evaluate the responses of Quercus crispula and Quercus dentata to herbivory, their leaves were subjected to simulated herbivory in early spring and examined for the subsequent changes in leaf traits and attacks by chewing herbivores in mid summer. In Quercus crispula, nitrogen content per area was higher in artificially damaged leaves than in control leaves. This species is assumed to increase the photosynthetic rate per area by increasing nitrogen content per area to compensate leaf area loss. In Quercus dentata, nitrogen content per area did not differ between artificially damaged and control leaves, while nitrogen content per mass was slightly lower in artificially damaged leaves. The difference in their responses can be attributable to the difference in the architecture of their leaves and/or the severeness of herbivory. The development of leaf area from early spring to mid summer was larger in artificially damaged leaves than in control leaves in both species, suggesting the compensatory response to leaf area loss. Leaf dry mass per unit area was also larger in artificially damaged leaves in both species, but the adaptive significance of this change is not clear. In spite of such changes in leaf traits, no difference was detected in the degree of damage by chewing herbivores between artificially damaged and controlled leaves in both species.     

Cassava Response to Water Deficit in Deep Pots: Root and Shoot Growth, ABA, and Carbohydrate Reserves in Stems, Leaves and Storage Roots

Cassava (Manihot esculenta, Crantz) is an important staple crop for tropical climates worldwide, including drought-prone environments where it is valued for its reliable yield. The extent to which stress tolerance involves regulation of growth and carbon balance aided by remobilization of carbohydrate from various plant parts was investigated. Plants were grown in 1-meter high pots to permit observation of deep rooting while they were subjected to four soil water regimes over a 30-d period. Transpiration declined abruptly in conjunction with leaf ABA accumulation and severe leaf abscission. In water stressed plants, growth of all plant parts decreased substantially; however, a basal rate of leaf growth continued to provide some new leaves, and although growth of fibrous lateral roots was reduced, main root elongation to deeper regions was only modestly decreased by stress. In leaf blades and petioles, sugars were the predominant form of nonstructural carbohydrate and about one third was in starch; these reserves were depleted rapidly during stress. In contrast, stems and storage roots maintained relatively high starch concentrations and contents per organ until final harvest. Stems gradually lost starch and had sufficient reserves to serve as a prolonged source of remobilized carbohydrate during stress. The amount of starch stored in stems represented about 35 % of the reserve carbohydrate in the plant at the onset of water stress (T₀), and 6 % of total plant dry mass. We suggest that this pool of carbohydrate reserves is important in sustaining meristems, growing organs, and respiring organs during a prolonged stress and providing reserves for regrowth upon resumed rainfall.     

CHANGES IN STARCH DISTRIBUTION IN THE OVERWINTERING ORGANS OF TYPHA LATIFOLIA (TYPHACEAE)

Histochemical determinations for storage of carbohydrates in rhizomes, roots, and young shoots of Typha latifolia L. (Typhaceae) were conducted during the overwintering period from November to April. Early winter analysis showed that rhizomes and roots contained large amounts of starch (45.03% and 22.80% dry weight, respectively). The major storage tissue was parenchyma of the rhizome central core. From winter into spring a gradual decrease in storage starch in the rhizome and root occurred concurrently with starch accumulation near zones of rapid development in young shoots (buds), but the rhizome retained much starch (27.40% dry weight) into the start of its 2nd yr.     

Foliage nitrogen turnover: differences among nitrogen absorbed at different times by Quercus serrata saplings

Background and Aims

Nitrogen turnover within plants has been intensively studied to better understand nitrogen use strategies. However, differences among the nitrogen absorbed at different times are not completely understood and the fate of nitrogen absorbed during winter is largely uncharacterized. In the present study, nitrogen absorbed at different times of the year (growing season, winter and previous growing season) was traced, and the within-leaf nitrogen turnover of a temperate deciduous oak Quercus serrata was investigated.

Methods

The contributions of nitrogen absorbed at the three different times to leaf construction, translocation during the growing season, and the leaf-level resorption efficiency during leaf senescence were compared using ¹⁵N.

Key Results

Winter- and previous growing season-absorbed nitrogen significantly contributed to leaf construction, although the contribution was smaller than that of growing season-absorbed nitrogen. On the other hand, the leaf-level resorption efficiency of winter- and previous growing season-absorbed nitrogen was higher than that of growing season-absorbed nitrogen, suggesting that older nitrogen is better retained in leaves than recently absorbed nitrogen.

Conclusions

The results demonstrate that nitrogen turnover in leaves varies with nitrogen absorption times. These findings are important for understanding plant nitrogen use strategies and nitrogen cycles in forest ecosystems.     

Mass loss,fungal colonisation and nutrient dynamics of Phragmites australis leaves during senescence and early aerial decay

This study examined the mass loss, fungal biomass, and nutrient dynamics of standing Phragmites australis leaf blades during senescence and early decay in littoral reed stands of two hardwater lakes. Green living leaves were tagged at defined canopy heights in early autumn (late August or early September) and periodically collected until all leaf blades had fallen off the parent shoot. Samples were analysed for leaf dry mass remaining, fungal biomass associated with leaves (ergosterol concentrations), and nitrogen and phosphorus concentrations. Considerable mass loss of leaves occurred in the standing position (up to 28%). Nitrogen and phosphorus concentrations of leaves decreased substantially with time (by 39–77%), indicating that a major portion of these nutrients was translocated to the rhizome during senescence. Fungal biomass associated with leaves increased during the study period, reaching an estimated maximum of about 40 mg g⁻¹ of leaf dry mass. Fungal biomass was negatively correlated with leaf N and P concentrations. The observed patterns of leaf mass loss, nutrient dynamics, and fungal biomass were consistent with the successive senescence and death of leaves from the shoot base to its tip. The results of this study point to a notable mass loss of P. australis leaf blades in the standing position, which appears to be mediated by both plant and microbial processes. Nutrient dynamics, in contrast, appear to be largely governed by plant processes.     

The effect of altered sink-source relations on photosynthetic traits and matter transport during the phase of reproductive growth in the annual herb <Emphasis Type="Italic">Chenopodium album</Emphasis>

Annual plants transport a large portion of carbohydrates and nitrogenous compounds from leaves to seeds during the phase of reproductive growth. This study aimed to clarify how reproductive growth affects photosynthetic traits in leaves and matter transport within the plant in the annual herb Chenopodium album L. Plants were grown in pots and either reproductive tissues or axillary leaves were removed at anthesis. Matter transport was evaluated as temporal changes in dry mass (as a substitute of carbohydrates) and nitrogen content of aboveground organs: leaves, axillary leaves, stems and reproductive tissues. Photosynthetic capacity (light-saturated photosynthetic rate under ambient CO₂ concentration), nitrogen, chlorophyll and soluble protein content were followed in the 20^th leaf that was mature at the start of the experiment. Removal of reproductive tissues resulted in accumulation of dry mass in leaves and axillary leaves, and accumulation of nitrogen in stem as nitrogen resorption from leaves and axillary leaves proceeded with time. Removal of axillary leaves proportionally reduced dry mass and nitrogen allocation to reproductive tissues, thus affecting the quantity but not quality of seeds. Removal treatments did not alter the time course of photosynthetic capacity, nitrogen, chlorophyll or soluble protein content during senescence in the 20^th leaf, but changed the photosynthetic capacity per unit of leaf nitrogen according to demand from reproductive tissues. Together, the results indicate that reproductive tissues affected carbon and nitrogen economy separately. The amount of carbon was adjusted in leaves through photosynthetic capacity and carbohydrate export from them, and the amount of nitrogen was adjusted by transport from stem to reproductive tissues. The plant’s ability to independently regulate carbon and nitrogen economy should be important in natural habitats where the plant carbon-nitrogen balance can easily be disturbed by external factors.     

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.     

The importance of nitrogen and carbohydrate storage for plant growth of the alpine herb Veratrum album

We examined whether nitrogen (N) and carbohydrates reserves allow Veratrum album, an alpine forb, to start spring growth earlier than the neighbouring vegetation and to survive unpredictable disturbances resulting in loss of above-ground biomass. * Seasonal dynamics of plant reserves, soil N availability and vegetation growth were monitored. Veratrum album shoots were experimentally removed when carbohydrate reserves were at a seasonal minimum and the subsequent changes in biomass and reserves were compared with those in control plants. Reserves did not give V. album a competitive advantage in spring; however, they did function as a buffer against the impact of calamities. Shoot removal resulted in significantly lower root dry weight, higher N concentration in rhizome and roots and lower starch concentrations in rhizome and roots but no plant mortality was observed. Veratrum album used stored N reserves to supplement N uptake and establish high leaf N concentrations, which facilitated a rapid refilling of depleted carbohydrate reserves. The primary function of N reserves appears to be to allow V. album to complete the growing cycle in as short a period as possible, thus minimizing exposure to above-ground risks.     

Fruit Development in Trillium : Dependence on Stem Carbohydrate Reserves

Leaves are the main source of carbon for fruit maturation in most species. However, in plants seeing contrasting light conditions such as some spring plants, carbon fixed during the spring could be used to support fruit development in the summer, when photosynthetic rates are low. We monitored carbohydrate content in the rhizome (a perennating organ) and the aboveground stem of trillium (Trillium erectum) over the entire growing season (May–November). At the beginning of the fruiting stage, stems carrying a developing fruit were harvested, their leaves were removed, and the leafless stems were maintained in aqueous solution under controlled conditions up to full fruit maturation. These experiments showed that stem carbohydrate content was sufficient to support fruit development in the absence of leaves and rhizome. This is the first reported case, to our knowledge, of complete fruit development sustained only by a temporary carbohydrate reservoir. This carbohydrate accumulation in the stem during the spring enables the plant to make better use of the high irradiances occurring at that time. Many other species might establish short-term carbohydrate reservoirs in response to seasonal changes in growing conditions.     

Devlopmental Physiology of Sugar-Beet: II. EFFECTS OF TEMPERATURE AND NITROGEN SUPPLY ON THE GROWTH, SOLUBLE CARBOHYDRATE CONTENT AND NITROGEN CONTENT OF LEAVES AND ROOTS

Plants were grown at temperatures of 15 and 25 ?C with two ratesof nitrogen supply. The changes in dry weight, leaf area, cellnumber, mean cell volume, soluble carbohydrate, and total nitrogenconcentration of the cotyledons, the first and second pair oftrue leaves, and the storage root were measured. Changes incell number and cell volume of the first pair of true leavesand storage root of plants were also measured at 11, 18, 25,and 32 ?C. Leaf growth before unfolding was chiefly by increase in cellnumber and after unfolding by increase in mean cell volume,while the growth of the storage root was almost entirely byincrease in cell number. The rates of cell division and cellexpansion were fastest at 25 ?C, but the initially high ratesof cell division in the terminal bud and in individual leavesdecreased rapidly and greater rates were maintained at the sub-optimaltemperatures, i.e. 15 and 18 ?C. After an initial period ofslow growth, the first-formed leaves grew faster and becamelarger at 15 than at 25 ?C. Leaves were produced, unfolded,grew faster, and became larger with increase in the externalconcentration of nitrogen, because cells divided and expandedfaster, so that nitrogen increased the number and size of cells. Sugar concentration was greater at 15 than at 25 ?C in leavesbut not in the storage root. Sugar concentration in the petiolesof the first and second pair of true leaves increased to 1.2and 2.0 per cent fresh weight respectively. Decreased nitrogensupply temporarily increased the sugar concentration of cotyledonpetioles and the seedling hypocotyl, but later decreased itin the leaves and storage root. Nitrogen concentration was greaterin the leaves and storage root at 15 than at 25 ?C with thelarger nitrogen supply. Nitrogen concentrations were similarin young leaves of all treatments but as the size of leavesincreased nitrogen concentrations decreased most rapidly at25 ?C with the smaller nitrogen supply. It is suggested that when increased leaf production and storage-rootgrowth occurs at temperatures below the growth optimum (25 ?C),they may be due to an effect of increased carbohydrate supplyon cell division and sugar storage.     

Leaf Construction Cost of the Most Abundant Species in an Upland Grassland Area of Northern Greece

The leaf construction cost, i.e., the energy expenditure required for the production of plant biomass (CC, g glucose/g dry biomass), is considered to be a major determinant of species success in various habitats. Nitrogen, carbon, and mineral contents in leaves were used to measure leaf CC. The aboveground biomass was sampled from the most abundant plant species (Poa pratensis L., Lolium perenne L., Festuca valida (Uechtr.) Penzes, Trifolium repens L., Taraxacum officinale Weber ex Wigg, Plantago lanceolata L., and Achillea millefolium L.) during the 1997 growing season in an upland grassland dominated by C₃ species. Soil samplings were performed in parallel with leaf samplings in order to determine soil inorganic nitrogen. T. repens leaves had the highest nitrogen concentration; grasses had the highest carbon content, while the highest mineral content was observed in the leaves of the forb species. The highest leaf CC was calculated for the legume T. repens followed by the grass F. valida. The grass L. perenne had the cheapest leaves, since it had the lowest CC. A positive correlation between leaf CC and soil inorganic nitrogen was evident for grasses (P. pratensis, L. perenne, F. valida) and P. lanceolata.     

Nitrogen acquisition,net production and allometry of <Emphasis Type="Italic">Alnus fruticosa</Emphasis> at a young moraine in Koryto Glacier Valley,Kamchatka, Russian Far East

Alders (Alnus spp.) often dominate at nutrient-poor sites by symbiotic relations with atmospheric nitrogen-fixing bacteria. However, little is known about quantitative relationships between root nodule as a nitrogen acquisition organ and leaf as a carbon acquisition organ. To examine carbon allocation, nitrogen acquisition and net production in nutrient-poor conditions, we examined allocation patterns among organs of shrub Alnus fruticosa at a young 80-year-old moraine in Kamchatka. Slopes of double-log allometric equations were significantly smaller than 1.0 for the root mass, leaf mass and root nodule mass against stem mass, and for the root nodule mass against root mass, indicating that smaller individuals invested disproportionally more biomass into resource-acquiring leaf and root tissues than to supportive tissues compared to older individuals. The slope of allometric equation of root depth against stem height was 0.542, indicating that smaller/younger individuals allocate disproportionally more biomass into root length growth than stem height growth. On the contrary, the root nodule mass isometrically scaled to leaf mass. The whole-plant nitrogen content also isometrically scaled to root nodule mass, indicating that a certain ratio of nitrogen acquisition depended on root nodules, irrespective of plant size. Although the net production per plant increased with the increase in stem mass, the slope of the double-log regression was smaller than 1.0. On the contrary, the net production per plant isometrically increased with leaf mass, root nodule mass and leaf nitrogen content per plant. Since the leaf mass isometrically scaled to root nodule mass, growth of each individual occurred at the leaves and root nodules in a coordinated manner. It is suggested that their isometric increase contributes to the increase in net production per plant for A. fruticosa in nutrient-poor conditions.     

Leaf age and season influence the relationships between leaf nitrogen, leaf mass per area and photosynthesis in maple and oak trees

Abstract. Seasonal changes in photosynthesis, leaf nitrogen (N) contents and leaf mass per area (LMA) were observed over three growing seasons in open-grown sun-lit leaves of red maple ( Acer rubrum ), sugar maple ( A. sacchamm ) and northern pin oak ( Quereus ellipsoidalis ) trees in southern Wisconsin. Net photosynthesis and leaf N were highly linearly correlated on both mass and area bases within all species from late spring until leaf senescence in fall. Very early in the growing season leaves had high N concentrations, but low photosynthetic rates per unit leaf N, suggesting that leaves were not fully functionally developed at that time. Leaf N per unit area and LMA had nonparallel seasonal patterns, resulting in differing relationships between leaf N/area and LMA in the "early versus late growing season. As a result of differences in seasonal patterns between leaf N/area and LMA, net photosynthesis/area was higher for a given LMA in the spring than fall, and the overall relationships between these two parameters were poor.     

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

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