玉米冠层内太阳直接辐射三维空间分布的模拟

太阳直接辐射在植物冠层内的空间分布特征影响植物生理生态功能 ,是衡量植物群体结构是否合理的重要指标。利用田间实测的玉米冠层内植株各器官的三维空间坐标进行冠层结构分析 ,将冠层内的植株器官表面划分成小面元 ;根据几何光学中光的直线传播原理 ,利用面元沿太阳光线的平行投影和投影深度排序 (Z- buffer)算法计算冠层内面元受太阳光直接照射的情况 ,建立了太阳直接辐射在玉米冠层内三维空间分布的模拟模型。模型可计算出作物冠层内任选植株的器官表面或冠层内地面上的太阳直射光斑 (Sunflecks)分布 ,也可输出选定空间位置或范围上的太阳直接辐射的分布 ,同时可实现模拟结果的三维可视化。根据此模型的模拟结果可对太阳直接辐射在玉米冠层内的空间分布进行各种分析。利用玉米冠层内光斑的三维分布测定试验 ,在光合有效辐射 (PAR)波段对模型进行了检验。模型适用于任意三维结构可测并可进行面元化划分的植物群体或个体     

Daylight spectra (400–740 nm) beneath sunny,blue skies in Tasmania,and the effect of a forest canopy

Abstract The photosynthetically active radiation (PAR) incident on a horizontal surface at an open mountain site is positively correlated with solar altitude for sunny, blue sky conditions. The proportion of red light in PAR decreases with increasing solar altitude, while that of blue increases. These results are consistent with the wavelength dependency of Rayleigh and Mie scattering. The ratio of near infrared radiation to PAR decreases with increasing solar altitude towards solar noon and with decreasing solar altitude towards sunset. Thus surface reflection seems to be an important part of the light climate. The relative transmission of daylight through a forest canopy to a horizontal surface is not correlated with solar altitude for sunny, blue sky conditions at a mountain site. The amount of diffuse daylight is negatively correlated with per cent canopy interception, and the amount of direct sunlight is negatively correlated with per cent solar track interception. Daylength is negatively correlated with both canopy and solar track interceptions. The proportion of red light in PAR increases with increasing solar altitude, while that of blue decreases. These results are opposite those for the open site and are due to the spatial patterns of canopy obstruction of the sky vault, and of the spectral quality of daylight across the sky. The ratio of near infrared radiation to PAR in shadelight increases with increasing canopy interception due to the selective scattering properties of the canopy. The ratio for shadelight is positively correlated with the ratio for sunflecks.     

Solar ultraviolet-B and photosynthetically active irradiance in the urban sub-canopy: A survey of influences

Stratospheric ozone loss in mid-latitudes is expected to increase the ultraviolet-B (UVB) radiation at the earth's surface. Impacts of this expected increase will depend on many factors, including the distribution of light in other wavelengths. Measurements of the photosynthetically active radiation (PAR) and UVB irradiance were made under clear skies at an open field and under the canopy of scattered trees in a suburban area in W. Lafayette, Indiana, USA (latitude 40.5°). Results showed that when there was significant sky view, the UVB penetration into sub-canopy spaces differs greatly from that of PAR. The UVBT _canopy (transmittance; irradiance below canopy/irradiance in open) was inversely related to sky view. The UVB irradiance did not vary as greatly between shaded and sunlit areas as did PAR. Analysis of measurements made near a brick wall indicated that the leaf area of a canopy and the brick wall primarily acted to block fractions of the sky radiance and contributed little scattered UVB to the horizontal plant. A model was developed to predict the UVB and PART _canopy based on diffuse fraction, sky view, and porosity of the crown(s) through which the beam is penetrating. The model accounted for the UVB and PART _canopy to within 0.13 and 0.05 root mean squared error (RMSE), respectively. Analysis of the errors due to model assumptions indicated that care must be taken in describing the sky radiance distribution, the porosity of trees, the penetration of diffuse radiation through porous trees, and the location of sky-obstructing trees and buildings.     

Carbon dioxide exchange and canopy conductance of two coniferous forests under various sky conditions

Sky conditions play an important role in the Earth’s climate system and CO₂ uptake by plants. We used eddy covariance and meteorological data, including global and diffuse photosynthetic photon flux density (PPFD), recorded over the 2008 and 2009 growing season at two Sitka spruce [Picea sitchensis (Bong.) Carr.] forest sites in northern Britain, in order to establish relationships between physiological properties under diverse sky conditions, i.e. (1) sunny, (2) cloudy, and (3) overcast, and several canopy activity-related properties. These properties are: (1) response to PPFD, (2) photosynthetic light use efficiency, and (3) canopy stomatal conductance. We found that Sitka spruce forests utilise PPFD in a more efficient way when solar radiation is dominated by diffuse radiation. Furthermore, our results show that diffuse radiation enhances canopy stomatal conductance, an effect which may be the result of both blue light enrichment within the canopy and the reduction in vapour pressure deficit during cloudy and overcast weather. Diffuse radiation does not only influence short-term (hourly, daily, monthly) canopy activity but also long-term forest growth.     

Leaf orientation and light interception by juvenile Pseudopanax crassifolius(Cunn.) C. Koch in a partially shaded forest environment

Leaf orientations and light environments were recorded for 40 juvenile Pseudopanax crassifolius trees growing in New Zealand in a partially shaded, secondary forest environment. Efficiencies of interception of diffuse and direct light by the observed leaf arrangments were calculated relative to those of three hypothetical leaf arrangements. Canopy gaps above the study plants were unevenly distributed with respect to azimuth and elevation above the horizon. Our results indicate that photosynthetically active radiation (PAR) received from the sides is more important than that received from directly above. In 33 of the plants leaf orientation was found to be significantly clustered towards one azimuth. The mean azimuth and the mean angle of declination were different for each plant. Leaves were steeply declined, and oriented towards the largest canopy gap at each site. Steep leaf angles reduced interception of direct and diffuse PAR when compared to interception by plant with a hypothetical horizontal leaf arrangement. When compared to a hypothetical arrangement with steep leaf declination and a uniform azimuth distribution, the observed leaf arrangement increased the efficiency of interception of diffuse PAR, but had a variable effect on the interception of direct PAR. Results indicate that the developing leaves of juvenile P. crassifolius orient towards the strongest sources of diffuse light, regardless of their value as a source of direct light. By maximising diffuse light interception while reducing direct light interception, leaf orientation may be a partial determinant of the types of habitats exploited by this species. This study emphasises the importance of considering diffuse light interception for plants growing in partially shaded environments.     

模拟降雨条件下玉米植株对降雨再分配过程的影响

为系统测定玉米(Zea mays)不同生长阶段的穿透雨、茎秆流和冠层截留,采用室内模拟降雨法测定了不同降雨强度、不同叶面积指数玉米冠下穿透雨和茎秆流,采用喷雾法测定了玉米不同生长阶段的冠层截留。对其进行了量化分析,并探讨了三者与玉米叶面积指数和降雨强度的关系,阐明了玉米冠下穿透雨的空间分布特征。结果表明:玉米冠下穿透雨量占冠上总降雨量比例为30.97%—94.02%,平均为63.92%;茎秆流量占降雨量比例的变化范围为5.68%—75.70%,平均为35.28%;冠层截留量在其全生育期内变化范围为0.02—0.43 mm,平均为0.16 mm,所占总降雨量比例最大仅为1%。随玉米生长,穿透雨量逐渐降低,茎秆流量和冠层截留量逐渐增加。穿透雨与茎秆流呈现此消彼长的关系,其中穿透雨率平均由93.55%降至36.23%;茎秆流率平均由5.98%增加至70.42%。降雨强度与穿透雨量和茎秆流量呈正相关关系,但是二者占总降雨量的比例与降雨强度关系不显著(P0.05)。随着玉米生长,穿透雨冠下空间分布由均匀逐渐趋向于不均匀,使降雨经过冠层后趋于向行中汇集,但在玉米生长后期,集中于行中的穿透雨量也因叶片衰败而随之降低。揭示了玉米对降雨的再分配作用特征,可为农田水分有效利用、农田生态水文过程机理和坡耕地土壤侵蚀防治提供理论依据。     

Some approaches for measuring and modelling riparian shade

The radiation environment of streams is of major ecological importance because it controls stream thermal regime and light availability for photosynthesis. Therefore, methods are needed for measuring stream shade in practical riparian management. The quantity ‘diffuse non-interceptance’ (difn), defined as the proportion of incident lighting received under a sky of uniform brightness and best estimated from fish-eye images, is useful for general specification of light exposure. For routine measurement of difn along stream reaches we recommend using a matched pair of simple light sensors (e.g. photosynthetically available radiation sensors) under conditions of complete overcast (which has almost uniform brightness). Methods are also needed for predicting future light exposure as riparian plantings grow and increasingly shade the stream. A simple model is outlined for predicting difn at the channel centre as a function of channel dimensions (stream width, w) and riparian plant character (foliage density, canopy height, h). The model reproduces the broad empirical trend of increasing shade with increasing h/w ratio. Future model refinement will aim to quantify the increase in shade moving from channel centre to edge under an overhanging canopy.     

Morphological responses of Datura ferox L. seedlings to the presence of neighbours

Summary We studied the effects of density on the dynamics of seedling growth and canopy microclimate within experimental stands composed of Datura ferox L. seedlings grown in individual pots. Interception of photosynthetically active radiation (PAR) by seedlings was evaluated either indirectly, by measuring leaf area, proportion of leaf area shaded by neighbouring individuals and laminar orientation with respect to sunlight, or directly, by measuring PAR at individual leaves at their natural angle of display. An integrating cylinder, with a geometry approximating that of a stem, was used within the canopies to measure the red:far-red (R:FR) ratio of the light flux from all compass points parallel to the soil surface. Seedlings responded rapidly (i.e. 1–2 weeks) to increased density by producing longer internodes and partitioning more dry matter to stems relative to leaves. These responses were observed before either PAR interception of growth rate were reduced by the presence of neighbours. Conversely, morphogenetic adjustment was preceded by a consistent effect of plant density on the R:FR ratio of the light received by the integrating cylinder. Air and soil temperature were not affected by density in these experiments. Differences in wind velocity within the canopy associated with plant density were avoided by the experimental procedure. The results support the idea that the drop in R:FR ratio of the light flux parallel to the ground — e.g. reflected sunlight — is an early signal that allows rapid adjustment of plant form to changes in canopy structure.     

Plant competition for light analyzed with a multispecies canopy model

Summary Competition for light among species in a mixed canopy can be assessed quantitatively by a simulation model which evaluates the importance of different morphological and photosynthetic characteristics of each species. A model was developed that simulates how the foliage of all species attenuate radiation in the canopy and how much radiation is received by foliage of each species. The model can account for different kinds of foliage (leaf blades, stems, etc.) for each species. The photosynthesis and transpiration for sunlit and shaded foliage of each species is also computed for different layers in the canopy. The model is an extension of previously described single-species canopy photosynthesis simulation models. Model predictions of the fraction of foliage sunlit and interception of light by sunlit and shaded foliage for monoculture and mixed canopies of wheat (Triticum aestivum) and wild oat (Avena fatua) in the field compared very well with measured values. The model was used to calculate light interception and canopy photosynthesis for both species of wheat/wild oat mixtures grown under normal solar and enhanced ultraviolet-B (290–320 nm) radiation (UV-B) in a glasshouse experiment with no root competition. In these experiments, measurements showed that the mixtures receiving enhanced UV-B radiation had a greater proportion of the total foliage area composed of wheat compared to mixtures in the control treatments. The difference in species foliage area and its position in the canopy resulted in a calculated increase in the portion of total canopy radiation interception and photosynthesis by wheat. This, in turn, is consistent with greater canopy biomass of wheat reported in canopies irradiated with supplemental UV-B.     

Measuring and modelling plant area index in beech stands

For some beech (Fagus sylvatica L.) stands with different stand densities the plant area index (PAI) was measured by means of a Licor LAI-2000 plant canopy analyser. The stands are located on the slopes of a valley in south-west Germany and had been treated by different types of silvicultural management (heavy shelterwood felling, light shelterwood felling, control plot). The analyser was used (a) to investigate the light conditions on plots of the same thinning regime, (b) to quantify the differences between the different treatments and (c) to obtain absolute values of PAI for interdisciplinary research. PAI was measured at three different phenological stages (leafless, leaf-unfolding and fully leafed season in 2000) and was found to be about 5.2 for the fully developed canopy on the control plots, 3.2 on the light fellings and about 2.0 for the heavy fellings. In the leafless period PAI was between 1.1 (control) and 0.4 (heavy felling). Measurements made in summer 2000 and summer 2002 were compared, and showed an increase of PAI, especially on the thinned plots. Measurements of photosynthetically active radiation (PAR) above and below the canopy in combination with measured PAI were used to apply Beers Law of radiation extinction to calculate the extinction coefficient k for different sky conditions and for the different growing seasons on the control plots. The extinction coefficient k for the beech stands was found to be between 0.99 and 1.39 in the leafless period, 0.62 to 0.91 during leaf unfolding and between 0.68 and 0.83 in the fully leafed period. Using PAR measurements and the k values obtained, the annual cycle of PAI was modelled inverting Beers Law.     

Can we use photography to estimate radiation interception by a crop canopy?

Accuracy of determining radiation interception, and hence radiation use efficiency, depends on the method of measuring photosynthetically active radiation intercepted. Methods vary, from expensive instruments such as Sunfleck ceptometers to simple methods such as digital photography. However, before universal use of digital photography there is need to determine its reliability and compare it with conventional, but expensive, methods. In a series of experiments at Lincoln, New Zealand, canopy development for barley, wheat, white clover and four forage brassica species was determined using both digital photographs and Sunfleck ceptometer. Values obtained were used to calculate conversion coefficient (Kf/Ki) ratios between the two methods. Digital photographs were taken at 45° and 90° for barley, wheat and white clover and at only 90° for brassicas. There was an interaction of effects of crop and cultivar for the cereal crops. Barley closed canopies earlier than wheat, and ‘Emir’ barley and ‘Stettler’ wheat had consistently higher canopy cover than ‘Golden Promise’ and ‘HY459′, respectively. Canopy cover was consistently larger at 45° than 90° for cereals. However, for white clover, the angle of digital photography was not important. There was also an interaction between effects of species and method of determining canopy cover for brassicas. Photographs gave higher cover values than ceptometer for forage rape and turnip, but the relationship was variable for forage kale and swede. Kf/Ki ratios of 1.0–1.10 for cereals, white clover and forage rape and turnip show that digital photographs can be used to estimated radiation interception, in place of Sunfleck ceptometer, for these crops.     

Spectral composition of photosynthetically active radiation penetrating into a Norway spruce canopy: the opposite dynamics of the blue/red spectral ratio during clear and overcast days

The spectral composition of photosynthetically active radiation (PAR) during clear and overcast days was studied above the canopy (U) and at two layers of a dense Norway spruce stand [Picea abies (L.) Karst.] characterized with an average LAI = 7.3 ± 0.8 (middle layer: M) and 12.3 ± 0.7 (lower layer: L). Whereas the spectral composition of PAR incoming on the canopy surface during cloudy days (characterized by diffuse index DI > 0.7) was almost independent of the solar elevation angle, the proportion of the blue-green spectral region of PAR was significantly reduced at low elevation angles during days with prevailing direct radiation (DI < 0.3). The PAR spectrum at both M and L levels was only slightly enriched in the green spectral region (more pronounced for DI < 0.3). The penetration of diffuse radiation into the canopy resulted in a slight (approx. 5%) reduction of the blue region proportion that remained stable during the day. On the contrary, under clear sky conditions the penetration of blue and red radiation was dependent on the solar elevation in an opposite manner in comparison with the spectral composition of PAR incident on canopy, giving almost twofold proportion of the blue part of the spectrum at a low elevation angle at M layer. We suggest that the blue enhancement of the spectrum within the Norway spruce canopy during clear days is due to a specific spatial arrangement of the assimilatory apparatus of a coniferous stand. Further, the possible consequences of the observed dynamics of the PAR spectrum inside the canopy during clear days on the efficiency of PAR absorption of the needles located within the canopy are discussed.     

Radiation frost susceptibility and the association between sky exposure and leaf size

Plants growing in exposed and sheltered habitats have characteristic leaf structure and physiology that are traditionally associated with the total amount of incident sunlight. However, greater sky exposure also increases the susceptibility of leaves to radiation frost. Plants with large horizontal broadleaves are particularly susceptible to both overheating during the day and freezing at night. Moreover, the combined effects of high daytime sun-exposure and nighttime frost susceptibility could be particularly stressful to plant tissues. The purpose of this study was to evaluate the influence of elevation and microsite exposure (i.e. net loss of longwave radiation) on frost susceptibility, as well as the corresponding intraspecific variation in leaf size in the subalpine daisy (Erigeron peregrinus). Measured decreases in upper hemisphere infrared radiation (sky IR) of 0.014 W m^-2 m^-1 occurred with increasing elevation, beyond decreases predicted due to changes in air temperature and water content, resulting in an average decrease of 0.029 W m^-2 m^-1. Previous equations of sky IR based on air temperature and humidity were improved by adding this elevational term (r ² improved from 0.52 to 0.71). In contrast, a mean decrease of 6.5 W m^-2 m^-1 occurred with increasing sky exposure across a subalpine meadow. Leaf size in Taraxacum officinale decreased linearly with increasing elevation and a corresponding decline in sky IR. No difference in daily solar irradiance was measured across the same elevational gradient. Also, E. peregrinus had smaller leaves at high elevation microsites with greater sky exposure and decreased sky IR, while there was a much weaker association between leaf size and the amount of total daily solar irradiance. Differences in plant leaf structure and physiology traditionally associated with daytime sun-exposure may also be influenced by nighttime sky exposure and the susceptibility to radiation frosts.     

The ratio of transmitted near-infrared radiation to photosynthetically active radiation (PAR) increases in proportion to the adsorbed PAR in the canopy

The daily total photosynthetically active radiation (400??00?nm, PAR) and near-infrared radiation (700??000?nm, NIR) were measured in the understory beneath the canopy (PARt and NIRt) and above the canopy (PARi and NIRi) of a Japanese cool-temperate deciduous broad-leaved forest during the snow-free period (May to November). The integration of spectral radiation for NIR and that for PAR, and the daily integrations of instantaneous NIR and PAR, reduced the noises from the optical difference in spectrum and from canopy structure heterogeneity, sky condition and solar elevation. PARi/PARt was linearly related to NIRt/PARt (R2?=?0.96). The effect of cloudiness was negligible, because the fluctuation of NIRi/PARi was quite small regardless of season and weather conditions compared with the range of NIRt/PARt in the forest. The ratio of NIRt/PARt beneath the canopy was log-linearly related to the in situ leaf area index (LAI) with a wide range from 0 to 5.25 (R2?=?0.97). We conclude that seasonal changes in fAPAR (=?1???PARt/PARi) and LAI of a canopy can be estimated with high accuracy by transmitted NIRt and PARt beneath the canopy.     

COMPARATIVE CACTUS ARCHITECTURE AND PAR INTERCEPTION

Because CO₂ uptake by cacti can be limited by low levels of photosynthetically active radiation (PAR) and because plant form affects PAR interception, various cactus forms were studied using a computer model, field measurements, and laboratory phototropic studies. Model predictions indicated that CO₂ uptake by individual stems at an equinox was greatest when the stems were vertical, but at the summer and the winter solstice CO₂ uptake was greatest for stems tilted 30° away from the equator. Stem tilting depended on form and taxonomic group; four barrel cacti in Ferocactus and in Copiapoa and four cylindropuntias in Opuntia tilted toward a horizontal light beam by an average of 18°, 48°, and 52°, respectively, after growth periods of 1 to 4 yr. In contrast, three columnar species showed no significant phototropic response, perhaps because structural stability requires their massive stems to be erect. Field plants of the dense, multiple-stemmed shrub Opuntia echinocarpa had stems which tended to radiate outward from the plant base, and, although this would not influence the total PAR intercepted, it would result in a more uniform PAR distribution and hence higher CO₂ uptake. For O. echinocarpa and the even denser, mound-forming Echinocereus engelmannii, PAR and chlorophyll decreased approximately exponentially with depth into the canopy. The canopies of O. echinocarpa and other cylindropuntias did not extend to the ground; in certain species, such truncation apparently resulted from a combination of very low PAR levels just below the lower canopy edge and the light-dependent growth responses of individual stems. In addition, although the canopy surfaces of O. echinocarpa and O. acanthocarpa tilted toward the equator by about 30°, the canopies of other cylindropuntias tilted less or not at all; the computer model predicted that a 30° tilt would decrease interstem shading, increase daily PAR, and increase nocturnal CO₂ uptake by as much as 54, 26, and 24%, respectively. Not only can the shape of cacti be affected by PAR, but also shape influences PAR interception and hence CO₂ uptake.     

Maize root system growth and development as influenced by phosphorus deficiency

Effects on leaf growth, biomass accumulation and root morphogenesis associated with the establishment of phosphorus (P) deficiency were studied on maize in order to test the hypothesis that the root system response can be accounted for by the effect of P deficiency on the carbon budget of the plant. P deprivation had a large and rapid negative effect on leaf expansion. For 7 d after P deprivation, the total dry matter production per plant was almost fully accounted for by the effect of P starvation on leaf growth and its subsequent effect on photosynthetically active radiation (PAR) interception. No strong effect of P deficiency was observed on the radiation use efficiency during this first period, although it was reduced thereafter. Root growth was slightly enhanced a few days after P starvation, but strongly reduced thereafter. The elongation rate of axile roots was maintained throughout the experiment, whereas emergence of new axile roots and elongation of first-order laterals were drastically reduced. The density of first-order laterals was not severely affected. These morphological responses are very similar to what is observed when root growth is limited by the availability in carbohydrates. The results are therefore compatible with the hypothesis that P deficiency mainly affects the root system morphology through its effect on the carbon budget of the plant with no additional specific effect of P deficiency on root morphogenesis. The drastic and early reduction of shoot growth after P deprivation may explain that more carbohydrates were available for root growth which was observed a few days after P starvation and reported by several authors. Later on, however, because of the reduced leaf area of P-deprived plants, their capacity to intercept light was severely reduced so that root growth was finally reduced.Keywords: Zea mays L., maize, phosphorus, root, root morphogenesis.      

Effects of ultraviolet–B irradiances on soybean

Soybean [Glycine max (L.) Merr. cv. Hardee] and wheat (Triticum aestivum L. cv. Jori) were grown from seed under four ultraviolet-B irradiances and four levels of photosynthetically active radiation in a factorial design. The effects of ultraviolet-B radiation on leaf number and area, total dry matter production, dry weight of component organs, and plant height were compared between soybean and wheat. Ultraviolet-B radiation effects were dependent upon the level of photosynthetically active radiation incident during growth. Wheat and soybeans were both affected by low ultraviolet-B radiation flux densities; however, they differed markedly in their growth responses and biomass allocation patterns. Substantial interactions between ultraviolet-B and photosynthetically active radiation indicate a need for the measurement of longer wavelength radiation when evaluating the effects of ultraviolet-B radiation on plant growth in natural conditions.     

Inversion of net ecosystem CO2 flux measurements for estimation of canopy PAR absorption

The fractional absorption of photosynthetically active radiation (f_PAR) is frequently a key variable in models describing terrestrial ecosystem–atmosphere interactions, carbon uptake, growth and biogeochemistry. We present a novel approach to the estimation of the fraction of incident photosynthetically active radiation absorbed by the photosynthetic components of a plant canopy (f_Chl). The method uses micrometeorological measurements of CO₂ flux and incident radiation to estimate light response parameters from which canopy structure is deduced. Data from two Ameriflux sites in Oklahoma, a tallgrass prairie site and a wheat site, are used to derive 7‐day moving average estimates of f_Chl during three years (1997–1999). The inverse estimates are compared to long‐term field measurements of PAR absorption. Good correlations are obtained when the field‐measured f_PAR is scaled by an estimate of the green fraction of total leaf area, although the inverse technique tends to be lower in value than the field measurements. The inverse estimates of f_Chl using CO₂ flux measurements are different from measurements of f_PAR that might be made by other, more direct, techniques. However, because the inverse estimates are based on observed canopy CO₂ uptake, they might be considered more biologically relevant than direct measurements that are affected by non‐physiologically active components of the canopy. With the increasing number of eddy covariance sites around the world the technique provides the opportunity to examine seasonal and inter‐annual variation in canopy structure and light harvesting capacity at individual sites. Furthermore, the inverse f_Chl provide a new source of data for development and testing of f_PAR retrieval using remote sensing. New remote sensing algorithms, or adjustments to existing algorithms, might thus become better conditioned to ‘biologically significant’ light absorption than currently possible.     

Resource availability at three presumed stages of a heathland succession on the Lüneburger Heide,Germany

Abstract. The availability of maj or plant resources was investigated in three vegetation types that were assumed to represent different stages of a secondary succession on heathland on the Lüneburger Heide, northwestern Germany. Canopy transmission and absorption of photosynthetically active radiation (PAR), soil-water availability, and nutrient (Ca, K, Mg, N, P) availability were monitored with high spatial and temporal resolution in (1) a Calluna vulgaris heathland, (2) a pioneer birch-pine forest and (3) a late-successional oak-beech forest, situated close to each other on comparable geological substrate (diluvial). Mean fractional transmission of PAR during summer decreased from 0.48 in the heathland to 0.04 in the oak-beech forest while the fractional canopy absorption increased from 0.49 to 0.92. Soil-water availability as indicated by the soil-water potential, was significantly influenced by differential canopy interception loss and characteristic rooting patterns in the three vegetation types. Annual mean nutrient concentrations in the equilibrium soil solution were similar or, for some elements, increased from the heathland to the birch-pine and the oak-beech forest despite a growing demand. A marked increase was found for the total nutrient pools in the soil-organic layer between early and late successional vegetation types. On the poor glacial parent material, nutrient pools seem to be strongly dependent on stand productivity and litter production which increased with succession. Thus, for nutrients, facilitation seems to be important in this type of succession.     

BRANCHING PATTERNS OF COLUMNAR CACTI: INFLUENCES ON PAR INTERCEPTION AND CO2 UPTAKE

Field measurements and a computer model were used to determine how stem shape and arrangement of stems in space affect interception of photosynthetically active radiation (PAR) and CO₂ uptake under otherwise optimal conditions for four species of columnar cacti (Carnegiea gigantea, Lophocereus schottii, Pachycereus pringlei, and Stenocereus thurberi). In simulations where the number of widely spaced stems was increased from 1 to 19 but plant volume remained constant, surface area and PAR interception increased, leading to 3-fold increases in whole-plant CO₂ uptake. Increasing the distance between stems from 0 cm to infinity decreased self-shading and increased predicted CO₂ uptake 4-fold. Stem length, diam, ribbing characteristics, and spine coverage also influenced PAR interception. The model indicated that the observed higher frequency of branches on the south side of the trunk of C. gigantea had only a slight, though positive, effect on CO₂ uptake for single-branched plants. Because of its greater surface area (A), a five-stemmed plant of C. gigantea typical for a field site near Tucson, Arizona was predicted to have 52% more CO₂ uptake than a single-stemmed plant of the same volume (V). Although large A/V decreases water storage per unit transpiring area, whole-plant CO₂ uptake can be increased when A/V is increased by branching for these constant-volume plants. However, the stems must be arranged to avoid excessive self-shading and thus keep the area below PAR compensation small.