Studies on the life history of an evergreen herb,Pyrola japonica,population on a forest floor in a warm temperate region

The life history and matter economy were studied on an evergreen herb,Pyrola japonica Klenze, population growing on the floor of a deciduous forest in a warm temperate region, central Japan. Seasonal changes in standing crop, bulk density and reserve substance of each organ and in leaf area were investigated over a year. Monthly and annual net production of the population were estimated based on the growth in dry weight and the seasonal dynamics of reserve substance. Seasonal peak value of the monthly net production was 16.7 g d.w.m⁻² in May. The annual net production was estimated to be 60.2 g d.w.m⁻². Two phases were recognized in the annual pattern of development and matter economy of this plant. Phase I, from April to June, was characterized by the development of aboveground vegetative organs and the consumption of reserve substance rrom old organs. Phase II, from July to March, was characterized by the development of reproductive and underground organs, and the accumulation of reserve substance. Phase II was further divided into two sub-phases according to the behavior of the reserve substance and the rate of net production. The production process of the population was compared with those of other evergreen herbs growing on the forest floor in warm temperate regions.     

Seasonal pattern of photosynthetic production in a subalpine evergreen herb,<Emphasis Type="Italic"> Pyrola incarnata</Emphasis>

The seasonal pattern of growth and matter production of Pyrola incarnata, an evergreen herb on the forest floor in subalpine deciduous forests, was analyzed to understand the ecological significance of evergreenness in a subalpine climate with a short growing season and low temperature. Net production was highest under favorable light conditions in spring after the disappearance of snow cover, and 68% of the annual net production was attained before the canopy tree foliage had fully expanded. Most of the photosynthetic production in this period was carried out with over-wintered leaves. This appears to be an advantage of evergreenness. New leaves and inflorescences had developed in the period. Positive net production was maintained under deteriorating light conditions during summer, when 32% of the annual net production occurred. This production was used mainly for growth of fruits and underground organs. The net production of P. incarnata during summer was much higher than that of a related species that inhabits warm-temperate regions, because of its higher photosynthetic activity rather than its lower respiratory losses. The storage of dry matter in leaves and underground organs was not conspicuous. Unlike the warm-temperate species and another subalpine species that inhabits higher altitudes, P. incarnata is not strongly dependent on its reserve matter for the development of new organs.     

Aboveground net production and dry matter allocation ofPinus pumila forests in the Kiso mountain range,central Japan

Aboveground net production rates of the subalpine stone pine (Pinus pumila) forests in central Japan were estimated by the summation method; net production was defined as the sum of annual biomass increment and annual loss due to death. In the two pine stands of different scrub heights, P1 (200 cm) and P2 (140 cm), aboveground biomass reached 177 and 126 ton ha⁻¹, respectively. Leaf biomass was about 14 ton ha⁻¹ in each stand. The estimates of aboveground net production during the 2 year period (1987–1989) averaged 4.1 and 3.7 ton ha⁻¹ y⁻¹ in P1 and P2, respectively, which were at the lowest among the pine forests in the world. Two indices of efficiency of energy fixation, that is, the ratio of net production to the total radiation during a growing season and the ratio of net production to total radiation per unit of leaf weight, were evaluated. Both efficiency indices for the twoP. pumila stands fell in the range obtained for other Japanese evergreen conifer forests. This suggested that the low annual net production of the stone pine stands were mainly due to a limitation in the length of the growing season. The pine forests were also characterized by a small allocation (about 17%) of aboveground net production into biomass increment, in comparison with other evergreen conifer forest types. Annual net carbon gain in theP. pumila forests was suggested to be largely invested in leaf production at the expense of the growth of woody parts.     

A quantitative study of the forest floor biomass,litter fall and nutrient return in a Pinus roxburghii forest in Kumaun Himalaya

The present paper reports on the forest floor biomass, litter fall, nutrient return and turnover of organic matter in a Pinus roxburghii forest in Kumaun Himalaya. Peak values of fresh leaf litter, partially decomposed litter and wood litter on the forest floor occurred in April, May and September, respectively. The relative contribution of partially decomposed material to total forest floor biomass remained greatest throughout the annual cycle. The biomass of herbaceous vegetation was maximal in September with a total annual net production of 151 g m^-2. The total annual litter fall was 895 g m^-2, of which tree, shrub and herb litters accounted for 82.4%, 0.6%, and 16.8%, respectively. Annual nutrient return in kg ha^-1 through litter fall amounted to 278.6 ash, 73.9 N, 5.5 P, 79.7 Ca, 15.1Mg, 20.7 K and 3.6Na. The turnover rate for tree litter was 48% and that for various nutrients on the forest floor ranged between 40–79%.     

Dry matter production, population structure and environmental conditions of the spring ephemeralErythronium japonicum growing in various habitats differing in sunlight exposure in cool temperate Japan

Phenology, dry matter production, population structure and environmental conditions were examined inErythronium japonicum Decne plants growing on the floor of a deciduous broad-leavedQuercus mongolica forest (Q.m. stand), an evergreen coniferousCryptomeria japonica plantation (C. j. stand) and bare ground left for 3 years after the clearing of a forest composed of youngQ. mongolica andPinus densiflora trees (bare stand) in the cool temperate zone of Japan. The average population density of the plants growing at theQ.m. stand was much higher than that observed at the bare stand, whereas the average number of flowering plants at the former stand was less than half of that at the latter. The population density and number of flowering plants growing at theC. j. stand were both less than 30% of those at theQ. m. stand. The number of seedlings at theQ. m. stand was much more than that at the bare andC. j. stands. Their survivorship rate over 1 year at the former stand also seems to be significantly higher than those at the other stands. Their aboveground and belowground parts at the bare stand were exposed to more severe heat and water stress than those at the other two stands. The net production per leaf area of the plants growing at theQ. m. stand was two and six times larger than those at the bare andC. j. stands, respectively. The plants at the bare stand did not use the available solar radiation as efficiently for dry matter production through photosynthesis as those growing at theQ. m. stand, whereas those at theC. j. stand are strongly restricted in their photosynthetic process by the significantly limited light condition on the floor of the evergreen coniferous plantation. The differences in the number of plants reaching sexual maturity, the density and structure of the population and the net production between their three habitats are discussed here from the standpoint of differences in environmental conditions.     

皖南黟县次生灌草丛生物量的研究

本文对地处中亚热带的安徽黟县次生灌丛和灌草丛以及常绿阔叶幼林共9个样地的生物量进行了研究。结果表明:黟县常绿阔叶幼林、次生灌丛和灌草丛的地上部分为81.74、16.83和5.28t/ha。所调查的5个群落类型的地上部分生物量和生产力可回归成线性方程Y=3.88+0.14X,净生产力随生物量的增加而提高。     

Temperature and substrate controls on intra-annual variation in ecosystem respiration in two subarctic vegetation types

Arctic ecosystems are important in the context of climate change because they are expected to undergo the most rapid temperature increases, and could provide a globally significant release of CO₂ to the atmosphere from their extensive bulk soil organic carbon reserves. Understanding the relative contributions of bulk soil organic matter and plant‐associated carbon pools to ecosystem respiration is critical to predicting the response of arctic ecosystem net carbon balance to climate change. In this study, we determined the variation in ecosystem respiration rates from birch forest understory and heath tundra vegetation types in northern Sweden through a full annual cycle. We used a plant biomass removal treatment to differentiate bulk soil organic matter respiration from total ecosystem respiration in each vegetation type. Plant‐associated and bulk soil organic matter carbon pools each contributed significantly to ecosystem respiration during most phases of winter and summer in the two vegetation types. Ecosystem respiration rates through the year did not differ significantly between vegetation types despite substantial differences in biomass pools, soil depth and temperature regime. Most (76–92%) of the intra‐annual variation in ecosystem respiration rates from these two common mesic subarctic ecosystems was explained using a first‐order exponential equation relating respiration to substrate chemical quality and soil temperature. Removal of plants and their current year's litter significantly reduced the sensitivity of ecosystem respiration to intra‐annual variations in soil temperature for both vegetation types, indicating that respiration derived from recent plant carbon fixation was more temperature sensitive than respiration from bulk soil organic matter carbon stores. Accurate assessment of the potential for positive feedbacks from high‐latitude ecosystems to CO₂‐induced climate change will require the development of ecosystem‐level physiological models of net carbon exchange that differentiate the responses of major C pools, that account for effects of vegetation type, and that integrate over summer and winter seasons.     

Developing a Multilayered Integrated Numerical Model of Surface Physics – Growing Plants Interaction (MINoSGI)

This paper describes a Multilayered Integrated Numerical Model of Surface Physics – Growing Plants Interaction (MINoSGI), which represents interactions between the dynamics of forest ecosystems and microclimate. Aiming at a large‐scale study in the future, we describe forest dynamics by using area‐averaged prognostic equations for thedistributions of plant density and plant weight with respect to plant height classes, instead of individual‐based treatments for small‐scale forest patches. Growth and mortality of plants are modelled based on the carbon balance of each plant height class. The area‐averaged microclimate (e.g., light, wind speed, temperature, humidity, CO₂ concentration) within the forest canopy is simulated by a one‐dimensional multilayer canopy model, which includes most of the physical and physiological processes that control the forest microclimate. Owing to its multilayered framework, a direct specification is possible for the difference in the growing environment among plants of different size and species. Given hourly meteorological conditions, the model outputs energy, water, CO₂ and momentum fluxes to and from a forest, of which the structure changes through competition among plants. The model's performance was tested by comparing its outputs with observed data on the development of plant size distribution taken over a 5‐year period in an evergreen coniferous (Cryptomeria japonica) forest. The model produced realistic estimates of the total biomass increments during the period. The ratio of net primary production to gross primary production (=0.45) was consistent with previous estimates for temperate forests. The bimodal seasonal pattern in net ecosystem production was similar to the seasonal trend in the CO₂ flux measured over a forest of the same species. Although some limitations due to the one‐dimensional representation of microclimate were noticeable, the model adequately simulated distributions of annual growth rate, plant weight and diameter across plant height classes. Since the basic equations can be extended to include the effect of spatial variability with marginal increase of computational costs, the present model framework is feasible for large‐scale studies.     

The nitrogen budget of a pine forest under free air CO2 enrichment

Elevated concentrations of atmospheric CO₂ increase plant biomass, net primary production (NPP) and plant demand for nitrogen (N). The demand for N set by rapid plant growth under elevated CO₂ could be met by increasing soil N availability or by greater efficiency of N uptake. Alternatively, plants could increase their nitrogen-use efficiency (NUE), thereby maintaining high rates of growth and NPP in the face of nutrient limitation. We quantified dry matter and N budgets for a young pine forest exposed to 4 years of elevated CO₂ using free-air CO₂ enrichment technology. We addressed three questions: Does elevated CO₂ increase forest NPP and the demand for N by vegetation? Is demand for N met by greater uptake from soils, a shift in the distribution of N between plants, microbes, and soils, or increases in NUE under elevated CO₂? Will soil N availability constrain the NPP response of this forest as CO₂ fumigation continues? A step-function increase in atmospheric CO₂ significantly increased NPP during the first 4 years of this study. Significant increases in NUE under elevated CO₂ modulated the average annual requirement for N by vegetation in the first and third growing seasons under elevated CO₂; the average stimulation of NPP in these years was 21% whereas the average annual stimulation of the N requirement was only 6%. In the second and fourth growing seasons, increases in NPP increased the annual requirement for N by 27-33%. Increases in the annual requirement for N were largely met by increases in N uptake from soils. Retranslocation of nutrients prior to senescence played only a minor role in supplying the additional N required by trees growing under elevated CO₂. NPP was highly correlated with between-plot variation in the annual rate of net N mineralization and CO₂ treatment. This demonstrates that NPP is co-limited by C availability, as CO₂ from the atmosphere, and N availability from soils. There is no evidence that soil N mineralization rates have increased under elevated CO₂. The correlation between NPP and N mineralization rates and the increase in the annual requirement for N in certain years imply that soil N availability may control the long-term productivity response of this ecosystem to elevated CO₂. Although we have no evidence suggesting that NPP is declining in response to >4 years of CO₂ fumigation, if the annual requirement of N continues to be stimulated by elevated CO₂, we predict that the productivity response of this forest ecosystem will decline over time.     

鼎湖山森林生态系统演替过程中的能量生态特征

以时空替代的方法,将灌草丛、针叶林、针阔叶混交林和季风常绿阔叶林等４个处于同一空间下的群落当作同一样落演替进程中的４个阶段,研究了鼎湖山南亚热带森林演替过程中的能量生态特征。结果表明,鼎湖山南亚热带森林群落演替过程中,其垂直层次、叶面积指数、冠层对太阳辐射能的截获量、叶生物量、总生物量、总初级生产力、总呼吸量、净初级生产力、枯树木现存量和年输入量、昆虫啃食量、群落的能量现存量等随演替的进程而增加,     

Studies on the life history of the population of an evergreen herb,Pyrola japonica,on the forest floor in a warm temperate region

Photosynthetic and respiratory activities and gross production in relation to temperature conditions were investigated in the population of an evergreen herb,Pyrola japonica, growing on the floor of a deciduous forest in the warm temperate region of central Japan. Analysis of the temperature-photosynthesis relationship ofP. japonica leaves during the growing season indicated distinct seasonal changes in the temperature optimum for photosynthesis. This population was found to be acclimatable to ambient air temperatures exceeding 15C, but this acclimation became less pronounced under thermal conditions below 15 C. This plant possessed narrow photosynthetic optima in the warm season but wide optima in the cold season. The shape of the temperature-respiration curve did not vary significantly with the months except for April. The Q₁₀ for respiration between 10 C and 20 C was calculated to be 1.93–2.65. Annual dry matter loss associated with respiration was estimated to amount to 159.1 g d.w.m⁻² based on the measurements of the seasonal changes in the respiratory activity of each organ. Gross production of this population was estimated to be 219.3 g d.w.m⁻² year⁻¹ as the sum total of the net production (60.2 g d.w.m⁻²year⁻¹) and the respiration. Monthly gross production was high in the early growing season, and low and stable in winter.     

云南哀牢山中山湿性常绿阔叶林生物量的初步研究

 中山湿性常绿阔叶林是云南省亚热带山地植被垂直带的主要类型,其中分布在哀牢山上的木果石栎、景东石栎、腾冲栲林(Lithocarpus xylocarpus、 L. chintungensis、Castanopsis wattii forest)尤其具有代表性。本文用收获法、相关曲线法测定并估算了该群落近熟林和成过熟林的生物量与年平均净积累量,结果分别为508.57t/hm2,12.1051t／(hm2·a);293.04t／hm2,7.7443t／(hm2·a),对产生差异的原因作了分析。林分的叶面积指繰果分别为508.57t/hm2,文中还提出了群落乔木优势种各器官生物量估测的回归模型,并从生物量的角度阐明建群种在群落中的地位。     

Production dependence of vegetative propagation inDisporum smilacinum A. Gray

The dependence of vegetative propagation on the production of individual plants was examined inDisporum smilacinum A. Gray on the basis of shading experiments and field surveys. This species typically showed four types of reproductive behavior: sterile plants producing one plantlet (no propagation), sterile plants producing more than two plantlets (vegetative propagation), fertile plants producing one plantlet (sexual reproduction) and fertile plants producing more than two plantlets (both sexual reproduction and vegetative propagation). The propagation ofD. smilacinum was clearly related to the annual net production of each individual plant. The probability of a mother plant producing more than two vegetative propagules (plantlets) increased with net production of the plant in the current year. The number of propagules per plant and runner length increased with net production. It was possible to explain the types of reproductive behavior of this species on the basis of both the initial plant size before sprouting and its net production during the growing season. There was a critical initial plant size for sexual reproduction and a critical level of production for vegetative propagation.     

Balance of Competitive and Facilitative Effects of Exotic Trees on a Native Patagonian Grass

The balance between facilitation and competition in plants changes with species characteristics and environmental conditions. Facilitative effects are common in natural ecosystems, particularly in stressful environments or years. Contrarily, in artificial associations of plants, such as agroforestry systems, some authors have suggested that even when facilitative effects may occur, net balance of tree effects on grasses is usually negative, particularly in dry environments. The aim of this study was to determine the net effect of the exotic ponderosa pine on the native grass Festuca pallescens (St. Ives) Parodi in agroforestry systems in Patagonia. Soil water content, plant water status, and relative growth were measured in the grass growing in different treatments (determined by tree cover level) during two growing seasons with contrasting climatic conditions. Facilitative effects of trees over grass water status were recorded only when water availability was high. A net negative effect was detected on dates when soil water content was very low and evaporative demand was high. The strength of these negative effects depended on tree density and climatic conditions, being higher in treatments with lower tree canopy cover. These results indicate that the positive effect of trees could only be expected under relatively low stress conditions. However, relative growth of grasses was always similar in plants growing in forested plots than in open grassland. Differences in biomass allocation for grasses growing in shade and open habitats may reconcile these contrary results. Our results highlight the importance of the physiology of a species (relative drought and shade tolerance) in determining the response of a plant to a particular interacting species.     

Photosynthesis and respiration in bladelets of Ecklonia cava Kjellman (Laminariales, Phaeophyta) in two localities with different temperature conditions

Characteristics of photosynthesis and respiration of bladelets were compared between Ecklonia cava Kjellman sporophytes growing in a warmer temperate locality (Tei, Kochi Pref., southern Japan) and in a cooler temperate locality (Nabeta, Shizuoka Pref., central Japan). Photosynthesis and respiration were measured with a differential gas-volumeter (Productmeter). In photosynthesis-light curves at 20°C, the rate of net photosynthesis was almost the same at light intensities lower than 25 μmol m⁻² s⁻¹ and the light-saturation occurred at 200–400 μmol m⁻²s⁻¹ in plants of both localities. The light-saturated net photosynthetic rates were higher in winter and spring than in summer and autumn in both plants. The optimum temperature for net photosynthesis at 400 μmol m⁻²s⁻¹ was 27°C throughout the year in the Tei plant and 25–27°C in the Nabeta plant. The decrease of net photosynthetic rates in the supraoptimal temperature range up to 29°C was sharper in winter and spring than in summer and autumn in both plants, being smaller in the Tei plant than in the Nabeta plant in all seasons. The dark respiration rate always increased with water temperature rise in both plants. No clear differences were found in the dark respiration rate between Tei and Nabeta plants except that when measured against dry weight, the Tei plant showed a slightly lower rate as compared with the Nabeta plant.     

Supply-side controls on soil respiration among Oregon forests

To test the hypothesis that variation in soil respiration is related to plant production across a diverse forested landscape, we compared annual soil respiration rates with net primary production and the subsequent allocation of carbon to various ecosystem pools, including leaves, fine roots, forests floor, and mineral soil for 36 independent plots arranged as three replicates of four age classes in three climatically distinct forest types. Across all plots, annual soil respiration was not correlated with aboveground net primary production (R²=0.06, P>0.1) but it was moderately correlated with belowground net primary production (R²=0.46, P<0.001). Despite the wide range in temperature and precipitation regimes experienced by these forests, all exhibited similar soil respiration per unit live fine root biomass, with about 5 g of carbon respired each year per 1 g of fine root carbon (R²=0.45, P<0.001). Annual soil respiration was only weakly correlated with dead carbon pools such as forest floor and mineral soil carbon (R²=0.14 and 0.12, respectively). Trends between soil respiration, production, and root mass among age classes within forest type were inconsistent and do not always reflect cross‐site trends. These results are consistent with a growing appreciation that soil respiration is strongly influenced by the supply of carbohydrates to roots and the rhizosphere, and that some regional patterns of soil respiration may depend more on belowground carbon allocation than the abiotic constraints imposed on subsequent metabolism.     

Water level drawdown makes boreal peatland vegetation more responsive to weather conditions

Climate warming and projected increase in summer droughts puts northern peatlands under pressure by subjecting them to a combination of gradual drying and extreme weather events. The combined effect of those on peatland functions is poorly known. Here, we studied the impact of long-term water level drawdown (WLD) and contrasting weather conditions on leaf phenology and biomass production of ground level vegetation in boreal peatlands. Data were collected during two contrasting growing seasons from a WLD experiment including a rich and a poor fen and an ombrotrophic bog. Results showed that WLD had a strong effect on both leaf area development and biomass production, and these responses differed between peatland types. In the poor fen and the bog, WLD increased plant growth, while in the rich fen, WLD reduced the growth of ground level vegetation. Plant groups differed in their response, as WLD reduced the growth of graminoids, while shrubs and tree seedlings benefited from it. In addition, the vegetation adjusted to the lower WTs, was more responsive to short-term climatic variations. The warmer summer resulted in a greater maximum and earlier peaking of leaf area index, and greater biomass production by vascular plants and Sphagnum mosses at WLD sites. In particular, graminoids benefitted from the warmer conditions. The change towards greater production in the WLD sites in general and during the warmer weather in particular, was related to the observed transition in plant functional type composition towards arboreal vegetation.     

基于森林资源清查资料的落叶松林生物量和净生长量估算模式

 丰富的森林资源清查资料是了解各类森林材积准确信息的重要途径,如果能将这些资源用于估算森林生物量和生产力的动态变化,不仅对于科学地指导森林的经营管理,而且对于全球变化的研究,特别是区域尺度的生产力模型验证,都具有重要意义。根据我国落叶松(Larix)林生物量和材积的实际调查资料,探讨了基于森林资源清查资料(森林材积V和林龄A)估算森林生物量和生产力的方法,指出无论是人工林还是天然林,落叶松林的生物量与其蓄积量、生产力与其年均净生物生产量(B/A)和年均净蓄积生产量(V/A)均呈双曲线关系,但落叶松林的生产力与其生物量(B)关系不明显,并分别建立了人工和天然落叶松林的相关模型;所建模型克服了将森林生物量与其蓄积量之比作为常数的不足,并考虑了林龄对于森林生产力的影响。     

Influence of floristic composition on the net primary production and dry matter turnover in a tropical grassland

Plant biomass, net primary productivity and dry matter turnover were studied in a grassland situated in a tropical monsoonal climate at Kurukshetra, India (29°58′N, 76°51′E). Based on differences in vegetation in response to microrelief, three stands were distinguished on the study site. The stand I was dominated by Sesbania bispinosa, stand II represented mixed grasses and stand III was dominated by Desmostachya bipinnata. Floristic composition of the three stands revealed the greatest number of species on stand II (75). The study of life form classes indicated a thero-cryptophytic flora. The biomass of live shoots in all the three stands attained a maximum value in September (424–1921 g m^-2) and below ground plant biomass in November (749–1868 g m^-2). The annual above ground net primary production was greatest on stand I (2143 g m^-2) and lowest on stand II (617 g m^-2). The rate of production was highest during the rainy season (15.34 to 3.18 g m^-2 day^-2). Below ground net production ranged from 1592 to 785 g m^-2 y^-2 and the rates were high in winter and summer seasons. Total annual net primary production was estimated to be 3141, 1403, 2493 and 2134 g m^-2 on stands I, II, III and on the grassland as a whole, respectively. The turnover of total plant biomass plus below ground biomass indicated almost a complete replacement of phytomass within the year. The system transfer functions showed greater transfer of material from total net primary production to the shoot compartment during rainy season and to the root compartment during winter and summer seasons.     

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.