Marinus J. A. Werger

Patterns of light and nitrogen distribution in relation to whole canopy carbon gain in C3 and C4 mono- and dicotyledonous species

An analytical model was used to describe the optimal nitrogen distribution. From this model, it was hypothesized that the non-uniformity of the nitrogen distribution increases with the canopy extinction rate for light and the total amount of free nitrogen in the canopy, and that it is independent of the slope of the relation between light saturated photosynthesis (Pm) and leaf nitrogen content (nL). These hypotheses were tested experimentally for plants with inherently different architectures and different photosynthetic modes. A garden experiment was carried out with a C3 monocot [rice, Oryza sativa (L.)], a C3 dicot [soybean, Glycine max (L.) Merr] a C4 monocot [sorghum, Sorghum bicolor (L.) Moensch] and a C4 dicot [amarantus, Amaranthus cruentus (L.)]. Leaf photosynthetic characteristics as well as light and nitrogen distribution in the canopies of dense stands of these species were measured. The dicot stands were found to have higher extinction coefficients for light than the monocot stands. Dicots also had more non-uniform N distribution patterns. The main difference between the C3 and C4 species was that the C4 species were found to have a greater slope value of the leaf-level Pm—nL relation. Patterns of N distribution were similar in stands of the C3 and C4 species. In general, these experimental results were in accordance with the model predictions, in that the pattern of nitrogen allocation in the canopy is mainly determined by the extinction coefficient for light and the total amount of free nitrogen.

Canopy structure and leaf nitrogen distribution in a stand of Lysimachia vulgaris L. as influenced by stand density

SummaryA hypothesis that a dense stand should develop a less uniform distribution of leaf nitrogen through the canopy than an open stand to increase total canopy photosynthesis was tested with experimentally established stands of Lysimachia vulgaris L. The effect of stand density on spatial variation of photon flux density, leaf nitrogen and specific leaf weight within the canopy was examined. Stand density had little effect on the value of the light extinction coefficient, but strongly affected the distribution of leaf nitrogen per unit area within a canopy. The open stand had more uniform distribution of leaf nitrogen than the dense stand. However, different light climates between stands explained only part of the variation of leaf nitrogen in the canopy. The specific leaf weight in the canopy increased with increasing relative photon flux density and with decreasing nitrogen concentration.

Canopy Structure and Photon Flux Partitioning Among Species in a Herbaceous Plant Community

This paper demonstrates a new analysis of photon flux partitioning among species and an evaluation of the efficiency of photon flux capturing in terms of biomass investment. For that purpose, distributions of aboveground biomass, leaf area, and photon flux density (PPFD) were determined with the stratified harvest method in a stand of a tall herbaceous community on a floating fen at the time of peak standing crop. The stand contained 11 species and the photon flux absorbed by each species in the stand was esti- mated. Three tall dominant species absorbed 75% of the incident PPFD, while eight short subordinate species absorbed 2.5%. Tall species in the canopy received higher PPFD av- eraged over leaf area (4)area). However, the PPFD absorbed per unit aboveground biomass (4)mass) of the tall species was not higher than that of the subordinate species. (Fmass is a product of the leaf area ratio (LAR, the ratio of leaf area to aboveground biomass) and

Light environment, sapling architecture, and leaf display in six rain forest tree species

Architecture and leaf display were compared in saplings of six rain forest tree species differing in shade tolerance. Saplings were selected along the whole light range encountered in a forest environment. Species differed largely in realized height and crown expansion per unit support biomass, but this could not be related to differences in shade tolerance. The results demonstrate that there exist various solutions to an effective expansion of plant height and crown area. It is argued that choice of the study species and the ontogenetic trajectory regarded determine to a large extent the outcome of interspecific comparisons. No evidence was found that pioneers were characterized by a multilayered and shade tolerants by a monolayered leaf distribution. Yet, sun plants had a similar crown area, a deeper crown, and a higher leaf area index compared to shade plants and their leaves were more evenly distributed along the stem. This suggests that differences in leaf layering are found between plants growing in different light environments, rather than between species differing in shade tolerance.

Tropical rain forest types and soil factors in a watershed area in Guyana

Abstract. An inventory was made of the vegetation and soils of a watershed area of 480 ha in the tropical rain forest of Guyana. 252 plots of 0.05 ha were sampled. In total 111 tree species > 20 cm were recorded. A TWINSPAN analysis resulted in seven groupings. Correspondence Analysis revealed that the major environmental differentiation underlying the floristic variation is according to soil type. Finally, five main forest communities were described. Most sites in the watershed area are characterized by the dominance of one or a few species. Species distribution patterns are strongly determined by soil type and drainage class.

Canopy Development and Leaf Nitrogen Distribution in a Stand of Carex Acutiformis

Seasonal changes in leaf nitrogen distribution were examined in the canopy of a Carex acutiformis stand in a wet meadow area. Although there was a tendency for leaf nitrogen concentration to decrease with increasing leaf age in any one layer of the canopy, nitrogen concentration increased significantly with plant height despite increasing age of leaf portions higher in the canopy. This suggests a predominant effect of the light climate on the nitrogen distribution within the canopy. During the growing period, standing crop dry mass increased significantly, while the increase in the standing crop of nitrogen was marginal. The amount of nitrogen decreased in the lower layers and increased in the upper layers, and a strongly decreasing gradient of nitrogen concentration developed from the top to the bottom of the canopy. It is suggested that this gradient resulted mainly from leaf tips with high nitrogen concentrations being lifted to higher positions because of growth at the base, with some retranslocation of nitrogen downwards from senescing tips to active parts. The distribution of nitrogen concentration became less uniform during the growing period, thus supporting the prediction that nitrogen concentration should become less uniformly distributed with development of the canopy.

Leaf nitrogen distribution and whole canopy photosynthetic carbon gain in herbaceous stands

The amount of photosynthetically-active photon flux density incident upon a leaf and the nitrogen content of that leaf strongly affect the photosynthetic carbon gain of that leaf. Therefore, the canopy structure of a stand, affecting the light climate in the canopy, and the leaf nitrogen distribution pattern in the canopy, affect the carbon gain of the whole canopy. This review discusses the results of studies directed to this problem and obtained so far in open and in dense canopies of stands of herbaceous, monocotyledonous or dicotyledonous, plants in their growing or flowering stages. It is found that the leaf nitrogen distribution pattern in the canopy is vertically non-uniform, and in dense stands more strongly so than in open stands. The leaf nitrogen distribution pattern in most canopies closely approaches an optimal pattern in that it maximizes whole canopy potential carbon gain as calculated for the actual total leaf nitrogen content and leaf area index of the stand. The resulting increase in potential carbon gain as compared to a uniform leaf nitrogen distribution pattern is considerable and it is larger in dense stands than in open stands. For at least some dense stands simulation studies show that with the available total leaf nitrogen content, whole canopy carbon gains could still be considerable higher had a lower leaf area index been developed.

The vertical distribution of nitrogen and photosynthetic activity at different plant densities in Carex acutiformis

Shoots of the monocotyledonous perennial Carex acutiformis were grown in open (28 shoots m−2) and dense stands (280 shoots m−2). For fully grown stands the distribution of relative PPFD and leaf nitrogen per unit leaf area over canopy depth was determined. Light response of photosynthesis was measured on leaf segments sampled at various heights in the stands. Relations between parameters of these curves and leaf nitrogen were investigated.Simulations showed that in the open stand daily canopy photosynthesis was not affected by nitrogen redistribution in the canopy. For the dense stand however, a uniform nitrogen distribution would lead to only 73% of the maximum net carbon gain by the stand under optimal nitrogen distribution. The actual canopy photosynthesis was only 7% less than this theoretical maximum; the actual nitrogen distribution of the dense stand clearly tended to the optimal distribution. The vertical pattern of the nitrogen distribution was to a large extent determined by the minimum leaf nitrogen content. The relatively high minimum leaf nitrogen content found for Carex leaves may perhaps be necessary to maintain the physiological function of the basal parts of the leaves.

Light partitioning among species and species replacement in early successional grasslands.

Abstract We studied canopy structure, shoot architecture and light harvesting efficiencies of the species (photon flux captured per unit above-ground plant mass) in a series of exclosures of different age (up to 4.5 yr) in originally heavily grazed grassland in N Japan. Vegetation height and Leaf Area Index (LAI) increased in the series and Zoysia japonica, the dominant in the beginning, was replaced by the much taller Miscanthus sinensis. We showed how this displacement in dominance can be explained by inherent constraints on the above-ground architecture of these two species. In all stands light capture of plants increased with their above-ground biomass but taller species were not necessarily more efficient in light harvesting. Some subordinate species grew disproportionally large leaf areas and persisted in the shady undergrowth. Some other species first grew taller and managed to stay in the better-lit parts of the canopy, but ultimately failed to match the height growth of their neighbours in this early successional series. Their light harvesting efficiencies declined and this probably led to their exclusion. By contrast, species that maintained their position high in the canopy managed to persist in the vegetation despite their relatively low light harvesting efficiencies. In the tallest stands ‘later successional’ species had higher light harvesting efficiencies for the same plant height than ‘early successional’ species which was mostly the result of the greater area to mass ratio (specific leaf area, SLA) of their leaves. This shows how plant stature, plasticity in above-ground biomass partitioning, and architectural constraints determine the ability of plants to efficiently capture light, which helps to explain species replacement in this early successional series. Nomenclature: Makino (1962); Ohwi (1965). Abbreviations: LAI = Leaf area index; LAR = Leaf Area Ratio; LMR = Leaf Mass Ratio; PPFD = Photosynthetically active photon flux density; SLA = Specific Leaf Area.

Photosynthetic capacity and nitrogen partitioning among species in the canopy of a herbaceous plant community

Partitioning of nitrogen among species was determined in a stand of a tall herbaceous community. Total amount of nitrogen in the aboveground biomass was 261 mmol N m−2, of which 92% was in three dominant species (Phragmites, Calamagrostis and Carex) and the rest was in the other eight subordinate species. Higher nitrogen concentrations per unit leaf area (nL) with increasing photosynthetically active photon flux density (PPFD) were observed in all species except for three short species. The changes in nL within species were mainly explained by the different nitrogen concentrations per unit leaf mass, while the differences in nL between species were explained by the different SLM (leaf mass per unit leaf area). Photon absorption per unit leaf nitrogen (ΦN) was determined for each species. If photosynthetic activity was proportional to photon absorption, ΦN should indicate in situ PNUE (photosynthetic nitrogen use efficiency). High ΦN of Calamagrostis (dominant) resulted from high photon absorption per unit leaf area (Φarea), whereas high ΦN of Scutellaria (subordinate) resulted from low nL although its Φarea was low. Species with cylinder-like “leaves” (Juncus and Equisetum) had low ΦN, which resulted from their high nL. Light-saturated CO2 exchange rates per unit leaf area (CER) and per unit leaf nitrogen (potential PNUE) were determined in seven species. Species with high CER and high nL (Phragmites, Carex and Juncus) had low potential PNUE, while species with low CER and low nL showed high potential PNUE. NUE (ratio of dry mass production to nitrogen uptake) was approximated as a reciprocal of plant nitrogen concentration. In most species, three measures of nitrogen use efficiency (NUE, ΦN and potential PNUE) showed strong conformity. Nitrogen use efficiency was high in Calamagrostis and Scutellaria, intermediate in Phragmites and relatively low in Carex. Nitrogen use efficiency of subordinate species was as high as or even higher than that of dominant species, which suggests that growth is co-limited by light and nitrogen in the subordinate species.