Canopy photosynthetic production in a Japanese larch forest. II. Estimation of the canopy photosynthetic production

Seasonal changes and yearly gross canopy photosynthetic production were estimated for an 18 year old Japanese larch (Larix leptolepis) forest between 1982 and 1984. A canopy photosynthesis model was applied for the estimation, which took into account the effect of light interception by the non-photosynthetic organs. Seasonal changes in photosynthetic ability, amount of canopy leaf area and light environment within the canopy were also taken into account. Amount of leaf area was estimated by the leaf area growth of a single leaf. The change of light environment within the canopy during the growing season was estimated with a light penetration model and the leaf increment within the canopy. Canopy respiration and surplus production were calculated as seasonal and yearly values for the three years studied. Mean yearly estimates of canopy photosynthesis, canopy respiration and surplus production were 37, 13 and 23 tCO₂ ha⁻¹ year⁻¹, respectively. Vertical trend, seasonal changes and yearly values of the estimates were analyzed in relation to environmental and stand factors.     

Crop row spacing and its influence on the partitioning of evapotranspiration by winter-grown wheat in Northern Syria

A study was conducted during the 1996–97 crop growth season at ICARDA in northern Syria, to investigate the influence of wheat canopy architecture on the partitioning of moisture between soil evaporation and crop transpiration, on a soil with high hydraulic conductivity. The study was conducted on the long-term two course wheat-lentil rotation trial, established on a swelling clay soil (Calcixerollic xerochrept). The wheat canopy architecture was manipulated by sowing the crop at either of two row-spacings, 0.17 or 0.30 m, both at a constant sowing rate equivalent to 120 kg ha^–1. In this study, evapotranspiration from the crop was inferred from changes in soil moisture content over time, evaporation and rainfall interception were measured daily using microlysimetry, drainage was estimated as being the difference between potential daily evapotranspiration, and the evapotranspiration estimated from the soil water deficit. Between sowing and day 80 (tillering stage), evapotranspiration was calculated to consist mainly of soil evaporation. However, after day 80, transpiration became an increasingly dominant component of evapotranspiration. For both row-spacings, cumulative evapotranspiration over the season was approximately 373 mm. In the narrow-row crop, transpiration and soil evaporation were approximately 185 mm and 183 mm of water respectively. Conversely for the wide row-spaced crop, 172 mm of water was transpired while about 205 mm of water evaporated from the soil surface. While green leaf area index did not differ between row-spacings, the architecture of the crops as a result of sowing affected solar radiation penetration such that more incident radiation was intercepted at the soil surface of the wide row-spaced crop. This is likely to have made some contribution to the elevated levels of evaporation from the soil beneath the canopy of the wide-sown crop.     

广东南岭天然常绿阔叶林林下光环境对林下幼树功能性状的影响

光环境与幼树功能性状的关系对天然林的更新与演替具有重要的生态学意义。以广东南岭区域天然常绿阔叶林下不同林龄(幼龄林,中龄林,老龄林)的森林群落为研究对象,通过监测冠层结构、林下光照数据和林下幼树功能性状等指标,研究林龄梯度下其冠层结构与林下光环境之间的关系,以及林下幼树功能性状对光环境的响应。结果表明:(1)中龄林叶面积指数显著高于幼龄林和老龄林(P0.05),随着林龄的增长,林冠开度和透光率逐渐下降,林龄梯度下透光率、R/FR(红光/远红光比值)、Bw/Rw(宽带蓝光/宽带红光比值)差异极显著(P0.001);(2)天然常绿阔叶林中透光率与光质之间极显著相关(P0.001),R/FR随着透光率的增加而增加,Bw/Rw随着透光率的增加而减少。(3)林下幼树功能性状在光环境之间差异显著(P0.05),老龄林林下幼树叶片氮含量显著高于幼龄林,而叶片重叠率显著低于幼龄林;(4)在本试验地中,R/FR和Bw/Rw的变化对林下幼树的高径比和光合作用并无显著影响,光强对同种植物不同光环境下最大净光合速率的影响较大。总体而言,林龄梯度冠层结构和光环境的差异能在一定程度上解释幼树功能性状的差异,这将有助于我们理解光环境对林下幼树更新的影响机制,同时为天然植被恢复和森林经营提供指导。     

亚热带常绿人工针叶林冠层内光能利用率

以我国南方主要造林树种湿地松(Pinus elliottii)、马尾松(P.massoniana)、杉木(Cunninghamia lanceolata)为研究对象,采用Li-6400便携式CO2/H2O红外气体分析仪测定生长中期(7月)和后期(10月)树冠内部不同层次(上层和下层)、方位(阳面和阴面)和叶龄(当年生和1年生)光环境、净光合速率(Pn),计算了光能利用效率(LUE)。结果表明:光合有效辐射(PAR)在树冠不同层次和方位间存在显著差异(P<0.05);当年生叶所处环境PAR高于1年生叶所处环境PAR,即在一定光环境之下发育的叶片往往被后来形成的新叶所遮荫。两时期3树种上层针叶的Pn均高于下层针叶;生长中期下层针叶的LUE高于上层叶,而生长后期层次间LUE相近或下层针叶不及上层针叶。不同方位针叶Pn因树种不同而不同,湿地松和马尾松阳面叶Pn高于阴面叶,而杉木阴面叶Pn高于阳面叶;两时期3树种阴面叶LUE均高于阳面叶。总体而言,叶片Pn和LUE对森林树冠内部光环境做出较强的可塑性。由于当年生针叶生理机能和光环境均优于1年生针叶,两时期3树种当年生针叶Pn均高于1年生针叶;生长中期湿地松和马尾松当年生...     

应用全天空照片估计林分透光孔隙度(郁闭度)

林分结构通过影响林内环境与生物因子,决定森林生态系统服务功能发挥;同时,通过调整林分结构亦可达到森林合理经营的目标。林分透光孔隙度(郁闭度)是决定林分结构的重要因子之一,它可以用来表征光、水等环境因子通过林冠进入林内的再分布状况。因此,合理、精确地确定林分内的郁闭状况,无论在森林经营与管理,还是在森林生态精细研究中,都显得十分重要。本文在以往研究的基础上,详细介绍了应用全天空照片(Hemispherical photograph/whole sky photograph)确定林分内一定高度(通常为1 m)的林分郁闭度或林分透光孔隙度(两者互补)的方法与步骤,重点讨论了在应用全天空照片确定林分郁闭度/透光孔隙度时选择的有效范围和应注意的实际问题。     

Implications of shoot structure on the rate of photosynthesis at different levels in a coniferous canopy using a model incorporating grouping and penumbra

1. The connection between high leaf area index (LAI) and photosynthetic production with two attributes of coniferous canopy structure: small leaf size and grouping of needles on shoots, was analysed using a simulation model.
2. The small size of conifer needles gives rise to penumbras, which even out the distribution of direct sunlight on the leaf area and thereby act to increase the rate of canopy photosynthesis per unit of LAI.
3. Grouping, by producing a non-uniform distribution of leaf area, causes a decrease in total canopy light interception at any given LAI, but improves the photosynthetic light capture by shoots in the lower canopy.
4. Application of the model on a case study showed that: (a) grouping had a negative effect on the rate of photosynthesis in the upper canopy, but deeper down in the canopy the situation was reversed; (b) in the lower canopy, photosynthetic rates were up to 10 times higher as a result from the combined effect of grouping and penumbra; (c) grouping did not improve the rate of canopy photosynthesis per unit of LAI, however, it can have a positive effect on the total photosynthetic production by allowing a higher productive LAI to be maintained; (d) penumbra, on the other hand, increased the rate of canopy photosynthesis by as much as 40% for moderate values of the LAI.     

Leaf area index (LAI) map of a protected area within the caldera of Vico Lake (Italy)

ABSTRACT

Structural traits of the vegetation types and plantations occurring in a protected area within the caldera of Vico Lake (Italy) were analysed. There were significant correlations among structural traits, at leaf and stand level. Leaf area index (LAI) and specific leaf area (SLA) were the most significantly correlated traits. LAI rose according to stand plant density, tree size and SLA; the highest LAI value monitored in the Fagus sylvatica L. forest was justified by the largest tree size (28.9±2.8 m height and 53±15 cm diameter) and the highest SLA (212±23 cm² g^-1). The main traits determining the variations in leaf structure among species were analysed by Principal Component Analysis (PCA). The LAI values were used to realise a map allowing us to delimit high LAI values (4.1–5.0), corresponding to the F. sylvatica forest and to the F. sylvatica forest with the sporadic presence of Quercus cerris L. and Castanea sativa Miller, mean LAI values (classes 3.1–4.0) corresponding to Corylus avellana L. plantations and to the Phragmites australis (Cav.) Trin. vegetation type, low LAI values (classes 2.6–3.0) corresponding to Q. cerris forests and C. sativa plantations.     

Reflection signals and the perception by phytochrome of the proximity of neighbouring vegetation

Abstract. Spectral photon distributions, red:far-red ratios (i.e. R:FR) and phytochrome photoequilibria (i.e. Pfr/Ptotal) were measured at various distances from artificial canopies composed of mustard or tobacco plants. Measurements were compared for radiation propagated predominantly vertically downwards and radiation propagated predominantly horizontally. Reflection signals from the artificial canopies were computed and shown to consist of a depletion of radiation over the 400–690 nm wavelength range, and an enhancement of radiation over the 690–800 nm range. R:FR and Pfr/Ptotal increased gradually with distance from the canopies, with significant depressions of both parameters evident at least as far as 30 cm from the vegetation stands. It is concluded that, in principle at least, detection of spectral quality differences by phytochrome would allow not only the presence but also the proximity of neighbouring plants to be perceived. Proximity perception is proposed as an ecologically valuable mechanism through which plants may be able to gauge their anticipatory responses to incipient shading according to the challenge posed by the nearness of neighbouring plants.     

Structural acclimation and radiation regime of silver fir (Abies alba Mill.) shoots along a light gradient

Shoot architecture has been investigated using the ratio of mean shoot silhouette area to total needle area ( ) as a structural index of needle clumping in shoot space, and as the effective extinction coefficient of needle area. Although can be used effectively for the prediction of canopy gap fraction, it does not provide information about the within‐shoot radiative regime. For this purpose, the estimation of three architectural properties of the shoots is required: needle area density, angular distribution and spatial aggregation. To estimate these features, we developed a method based on the inversion of a Markov three‐dimensional interception model. This approach is based on the turbid medium approximation for needle area in the shoot volume, and assumes an ellipsoidal angular distribution of the normals to the needle area. Observed shoot dimensions and silhouette areas for different vertical and azimuth angles (A_S) are used as model inputs. The shape coefficient of the ellipsoidal distribution (c) and the Markov clumping index (λ₀) are estimated by a least square procedure, in order to minimize the differences between model prediction and measurements of A_S. This methodology was applied to silver fir (Abies alba Mill.) shoots collected in a mixed fir–beech–spruce forest in the Italian Alps. The model worked effectively over the entire range of shoot morphologies: c ranged from 1 to 8 and λ₀ from 0·3 to 1 moving from the top to the base of the canopy. Finally, the shoot model was applied to reconstruct the within‐shoot light regime, and the potential of this technique in upscaling photosynthesis to the canopy level is discussed.     

Seasonal Direct Light Availability Affects Mean Leaf Orientation in a Herbaceous Multi-Species Canopy

Abstract: Relationship between leaf orientation in natural or semi-natural herbaceous communities and the local abundance of direct light has rarely been investigated. We present evidence from a permanent plot experiment that seasonal direct light availability in 40 × 40 cm plots significantly affects mean leaf orientation in a semi-natural wooded meadow. In sunny years and in well-illuminated conditions, the leaf surface is exposed predominantly to the north, and vice versa . Two groups of species are responsible for such a response: of the variation in leaf orientation, 41 % is explainable through direct light availability for light-demanding species, and 26 % for graminoid species. In shade-tolerant species and in forbs there is no sign of a regression between leaf orientation and direct light abundance. Ability to plastically react to variation in seasonal and local direct light availability appears to be a trait under considerable selective pressure only in light-demanding plants, and more characteristic of species with graminoid growth form.     

Ground monitoring the light–shadow windows of a tree canopy to yield canopy light interception and morphological traits

Monitoring the light–shadow windows of a tree via a grid system on the ground was performed on sunny summer days at high spatial resolution using a custom‐built, inexpensive scanner. The measurements were taken with two goals: (1) to quickly and remotely quantify the overall, short‐wave solar radiation (300–1100 nm) intercepted by the tree canopy, and (2) to yield such crown geometric traits as shape, size and the number of theoretical canopy leaf layers (leaf layer index, LLI) in relation to the section orthogonal to sunbeam direction (sun window). The ground readings at each measurement over the day were used to project a digitized shadow image. Image processing was applied and the intercepted radiation was calculated as the difference from the corresponding incoming radiation above the canopy. Tree‐crown size and shape were profiled via computer imaging by analysing the different shadow images acquired at the various solar positions during the day. It is notable that these combined images yielded the crown features without having to parameterize such canopy characteristics as foliage extension and spatial distribution.     

山西灵空山天然松栎混交林群落特征与冠层结构

以2011年建设的山西灵空山4 hm²天然松栎混交林森林动态监测样地为研究平台,以400个10 m×10 m样方为测量单元,于2016年进行群落特征研究,采用半球面影像法(DHP)分析冠层结构和林下光照特征.结果表明: 样地内共有乔木5558株,共计25种,分属于10科15属.冠层开阔度(CO)集中在15.0%～25.0%,叶面积指数(LAI)集中在1.5～2.5,林下光环境参数集中在10.0%～30.0%.建群种在样地内的分布对冠层结构和林下光环境影响显著;冠层结构对林下光环境所有参数的影响方向一致,其中采用叶面积指数评价冠层结构动态的效果更佳;冠层开阔度和叶面积指数对林下光环境产生相反的影响,且均对散射光入射率影响程度最大.温性松栎混交林的林冠层整体较为均匀,林下光分布较为集中,林分树种组成与冠层结构对林下光照影响显著.     

Light climate as a factor in the morphological variation of Atherosperma moschatum in a Tasmanian forest

Abstract The growth of saplings of Atherosperma moschatum within a Tasmanian forest during 1987/88 was negatively correlated with canopy closure. The dry weights and specific leaf weights of new leaves were also negatively correlated with canopy closure, but leaf areas showed a maximum at an intermediate canopy closure. Leaf chlorophyll concentration was positively correlated with canopy closure. Internode length was not correlated with canopy closure, but the ratio of leaf dry weight to internode length was negatively correlated with canopy closure. These results indicate that the load of photosynthetically active radiation may be a major determinant of the variation in plant development.     

Rapid assessment of forest canopy and light regime using smartphone hemispherical photography

Hemispherical photography (HP), implemented with cameras equipped with “fisheye” lenses, is a widely used method for describing forest canopies and light regimes. A promising technological advance is the availability of low‐cost fisheye lenses for smartphone cameras. However, smartphone camera sensors cannot record a full hemisphere. We investigate whether smartphone HP is a cheaper and faster but still adequate operational alternative to traditional cameras for describing forest canopies and light regimes. We collected hemispherical pictures with both smartphone and traditional cameras in 223 forest sample points, across different overstory species and canopy densities. The smartphone image acquisition followed a faster and simpler protocol than that for the traditional camera. We automatically thresholded all images. We processed the traditional camera images for Canopy Openness (CO) and Site Factor estimation. For smartphone images, we took two pictures with different orientations per point and used two processing protocols: (i) we estimated and averaged total canopy gap from the two single pictures, and (ii) merging the two pictures together, we formed images closer to full hemispheres and estimated from them CO and Site Factors. We compared the same parameters obtained from different cameras and estimated generalized linear mixed models (GLMMs) between them. Total canopy gap estimated from the first processing protocol for smartphone pictures was on average significantly higher than CO estimated from traditional camera images, although with a consistent bias. Canopy Openness and Site Factors estimated from merged smartphone pictures of the second processing protocol were on average significantly higher than those from traditional cameras images, although with relatively little absolute differences and scatter. Smartphone HP is an acceptable alternative to HP using traditional cameras, providing similar results with a faster and cheaper methodology. Smartphone outputs can be directly used as they are for ecological studies, or converted with specific models for a better comparison to traditional cameras.     

Understanding the intertidal zonation of two estuarine red macroalgae from ex situ photoacclimative responses

This study investigated the influence of light on the distributional limits of Bostrychia scorpioides and Catenella caespitosa within a low amplitude intertidal zonation in the Palmones estuary, Spain. Changes in photosynthesis- and growth-irradiance curves and pigment content were examined ex situ at irradiances found in their natural habitat (10–230 µmol photons m^?2 s^?1). The highest maximum photosynthesis rate (P_m) and photosynthetic efficiency (α) were found between 20–40 µmol photons m^?2 s^?1 in both species. Bostrychia scorpioides, the uppermost intertidal species, had a higher P_m, α, dark respiration rate and light compensation point (E_c) than C. caespitosa at all acclimation irradiances. Net photosynthetic rates measured at their respective acclimation irradiances showed photosynthetic responses of B. scorpioides to be maximized at high irradiances, while in C. caespitosa they did not change. Higher growth rates were obtained in C. caespitosa than in B. scorpioides, which may be related to its lower thallus specific carbon content. When irradiance decreased, the chlorophyll a content of B. scorpioides increased whereas in C. caespitosa R-phycoerythrin increased. E_c for growth of B. scorpioides coincided with the irradiance at its lower distributional limit in the estuary, below which this species showed losses in biomass. However, in C. caespitosa the sustained growth ex situ at saturating irradiances contrasts with its absence from the upper intertidal zone, where similar light regimes occur. We demonstrated that while light clearly restricts the growth of B. scorpioides to the uppermost intertidal zone, this environmental factor would not prevent C. caespitosa from growing at higher levels.     

林窗内光照强度的测量方法

林窗内光强存在复杂的时空变化,对植物更新和生长有着重要影响,因此,林窗光照强度的快速测量方法是生态学家十分关注的问题.目前,测量林窗光强的方法可分为3类:(1)直接测量法采用光量子探头等仪器直接测量光强,但测量林窗光强异质性时十分费时费力.(2)模型估测法通过几何计算可快速估测林窗任意位置光强,但模型估测法将林窗简化为圆柱体或椭圆柱体,并忽略了许多林窗光强的影响因素,这极大影响了它的测量精度.(3)相片法采用半球面影像等相片间接计算相片拍摄点的光强,但测量林窗光强异质性时需要在林窗内拍摄大量相片;相片法具有较高精度,可区分直射光和散射光,其中,基于半球面影像的林窗光指数(gap light index)精度最高,使用广泛;基于几何计算的林窗光指数不仅具有较高精度,且可以快速测量林窗任意位置光强,该方法适用于林窗光强水平分布格局、垂直结构以及光组成成分(直射光和散射光)特征研究.     

伏牛山自然保护区森林冠层结构对林下植被特征的影响

在伏牛山自然保护区典型地段设立样方,测定了森林生态系统内几种典型群落类型的冠层结构、光环境特征,调查了林下植被的特征,分析了它们之间的相互关系.结果显示:各群落的冠层结构和光环境有一定的差异,单因素方差分析表明,部分群落间的差异性达到显著水平;各群落灌木层物种丰富度、多样性和均匀度均高于草本层,而优势度正相反;线性拟合的结果表明,草本层的物种丰富度、多样性与冠下光合量子通量密度间呈极显著负相关,优势度与冠下光合量子通量密度间呈显著正相关,灌木层各参数与冠层结构特征间相关性不显著.研究表明,冠层结构的变化对草本层(包括更新幼苗)的影响显著高于灌木层.林隙/林窗或林中空地的出现可能对草本物种或其他阳性及先锋物种具有促进作用,而对优势种幼苗的萌发和定植产生负效应.推测在典型的落叶阔叶林生态系统演替进程中,林下光照强度可能不是最主要的限制因素,优势种种子的扩散、萌发和定植限制可能更重要.     

How tree species identity and diversity affect light transmittance to the understory in mature temperate forests

Light is a key resource for plant growth and is of particular importance in forest ecosystems, because of the strong vertical structure leading to successive light interception from canopy to forest floor. Tree species differ in the quantity and heterogeneity of light they transmit. We expect decreases in both the quantity and spatial heterogeneity of light transmittance in mixed stands relative to monocultures, due to complementarity effects and niche filling. We tested the degree to which tree species identity and diversity affected, via differences in tree and shrub cover, the spatiotemporal variation in light availability before, during, and after leaf expansion. Plots with different combinations of three tree species with contrasting light transmittance were selected to obtain a diversity gradient from monocultures to three species mixtures. Light transmittance to the forest floor was measured with hemispherical photography. Increased tree diversity led to increased canopy packing and decreased spatial light heterogeneity at the forest floor in all of the time periods. During leaf expansion, light transmittance did differ between the different tree species and timing of leaf expansion might thus be an important source of variation in light regimes for understory plant species. Although light transmittance at the canopy level after leaf expansion was not measured directly, it most likely differed between tree species and decreased in mixtures due to canopy packing. A complementary shrub layer led, however, to similar light levels at the forest floor in all species combinations in our plots. Synthesis. We find that a complementary shrub layer exploits the higher light availability in particular tree species combinations. Resources at the forest floor are thus ultimately determined by the combined effect of the tree and shrub layer. Mixing species led to less heterogeneity in the amount of light, reducing abiotic niche variability.     

Experimental effects of kelp canopies on subtidal coralline algae

Plants are often grouped as canopy species or understorey species because it is thought that that these sets of taxa interact in predictable ways. Mensurative experiments in southern Australia demonstrated that the percentage cover of encrusting coralline algae was greater, and articulated (branching) coralline algae less, on boulders under a canopy of dense kelp (>7 plants per m²), Ecklonia radiata, than on boulders without kelp. Experimental clearances of kelp and reciprocal transplants of boulders between patches of E. radiata and patches without kelp showed that canopies maintained and facilitated the growth of encrusting coralline algae and reduced the cover of articulated coralline algae. Potential artefacts associated with clearing kelp and transplanting boulders were not detected when tested with a series of translocation controls. These results reject the model that the co‐occurrence of E. radiata and encrusting corallines is just an assemblage of plants caused by spatial and temporal coincidence. Instead, they support the model that kelp facilitates the growth and survival of understorey algae.