间伐改形对陇东高原密闭富士苹果园冠层微域环境及叶片生理特性的影响

为了研究间伐改形对成龄乔化密闭红富士苹果园冠层微域环境、叶片显微结构、叶片生理特性和光合能力的影响,以16年生密闭红富士苹果园为研究对象,对果园冠层相对光照强度、温度、相对湿度、叶片叶绿素含量、显微结构、叶片光合和荧光等参数进行了测定。结果表明: 间伐改形后树体冠层相对光照强度、温度得到显著改善,分布更均衡,>30%的有效光强是对照(未间伐改形,CK)的1.57倍,温度比CK平均高1.1 ℃;间伐树体叶片叶绿素、叶片厚度、栅栏组织厚度显著提高,分别比CK提高了8.7%、5.4%、9.2%;叶片净光合速率、蒸腾速率、气孔导度也显著提高,分别比CK提高了12.6%、17.1%和7.3%。间伐果园和密闭果园叶片光合作用均受非气孔因素限制,间伐树体叶片的PSⅡ最大荧光产量和非光化学猝灭系数比CK提高了1.5%和2.1%。间伐改形后,叶片并未发生强光抑制,叶片单位反应中心吸收的光能、捕获的用于电子传递的能量和用于还原Q_A的能量得到显著提高。叶片生理特性与所处的光照、温度环境密切相关,密闭果园间伐改形后,果园冠层光照、温度得到改善,促进了叶片生长发育,改善了叶片显微结构,提高了叶片光合效能,是适宜陇东高原苹果产区密闭红富士果园调整和优化的关键措施。     

苹果密植园与间伐园树冠层内叶片光合潜力比较

通过对成龄苹果密植园和间伐园树冠不同层次和部位叶片光合潜力及辐射通量密度、叶片N含量和比叶重等指标的比较分析,研究了苹果园改造前后辐射能和氮素利用效率差异及其与产量品质的关系.结果表明：间伐显著改善了冠层内的辐射环境,间伐园冠层内的辐射分布明显比密植园均匀,相对辐射通量密度小于30%的无效光区接近0,而密植园冠层内的最低相对辐射通量密度为17%,在相对高度03以下均为无效光区;间伐园内冠层叶片的光合效率显著提高,间伐园树冠中、下部叶片的光合速率比密植园分别提高了78%和102%;叶片的最大羧化速率和最大电子传递速率也有较大幅度的提升.苹果园冠层叶片的光合效率与叶片N含量存在显著的相关关系,而叶片N含量又与辐射通量密度存在显著的相关关系,因此,可根据冠层叶片相对N含量的垂直分布间接和定量地判断叶片的光合效率或相对辐射通量密度的空间分布.     

间伐对杨桦次生林冠层结构及林下光照的影响

自然或人为干扰产生的森林冠层结构和林下光照差异是造成林下环境异质性、物种多样性的主要原因,对森林生态系统的结构、过程与格局具有重要作用.以桃山林场2块50 m×50 m的杨桦次生林固定样地为研究对象,分别作为间伐和对照样地,于2012—2016年、2018年获取样地冠层影像,通过方差分析、马尔科夫模型等方法分析其冠层结构、林下光照动态及间伐后林冠的恢复规律.结果表明: 间伐措施能显著调整冠层结构和林下光照,并能持续较长的作用时间,但调整效果随时间逐渐降低;间伐后冠层结构及林下光照的变化速度随时间推移而降低,间伐后前3年林冠恢复速度较快,3年后恢复速度降低并趋于平缓;林下光照与林冠开度呈显著正相关,与叶面积指数呈显著负相关,其中林下散射辐射与冠层结构参数的相关性最大,且间伐样地冠层结构与林下光照的相关性大于对照样地;间伐后冠层的恢复速度与林冠开度大小有关,林冠开度越大,林冠恢复越快,向更小林冠开度转移的时间越早.马尔科夫模型能够模拟冠层结构分布的变化,可用来预测冠层结构动态.     

不同土壤水分胁迫下沙漠葳的生长及生物量的分配特征

将从美国西部引进的2年生的沙生灌木沙漠葳（Chilopsis linearis）分别盆栽于含水量不同的土壤中,研究其生长及生物量的分配特征.结果表明,土壤水分胁迫严重限制了沙漠葳的营养生长和生殖生长,使单株叶片数、分枝数和侧根数显著下降,生物量大大降低,其中中度和重度胁迫下沙漠葳的干重分别比轻度胁迫降低40.9%和76.4%.重度土壤水分胁迫下沙漠葳的单叶干重、单叶面积和单位叶面积干重分别比轻度土壤水分胁迫降低63.45%、47.39%和27.23%,比叶面积和根茎比分别上升22.28%和86%.随土壤水分胁迫的加重,光合物质的积累从中下部叶片向中上部叶片转移.各构件生物量随土壤水分胁迫的加重而降低,其幅度大小为叶生物量＞茎生物量＞主根生物量＞侧根生物量,反应了沙漠葳对土壤水分胁迫响应的整体行动.     

不同树形龙安柚冠层特性

以开心形、Y字形、双层分层形和自然圆头形的龙安柚为试验材料,比较不同树形的冠层特性、叶片结构和生理特征,以期为龙安柚果园小环境的调控提供理论基础。结果表明:(1)开心形间隙分数阈值最高,是自然圆头形的4.33倍。开心形与Y字形的冠层光合辐射与透射系数均显著高于其他树形,但二者无显著差异,表明开心形和Y字形的冠层通风透光特性较好。(2)开心形与Y字形叶片厚度增加,叶面积和气孔密度较大,栅栏/海绵组织厚度和组织紧密度较高,叶片组织疏密度较低,且二者无显著差异,表明开心形与Y字形利于提高叶片的光合作用,降低蒸腾作用。(3)Y字形和开心形净光合速率、水分利用率、最大表观电子传递速率、初始斜率和半饱和光强较高,而蒸腾速率较低,二者对强光的耐受能力较强;其中开心形蒸腾速率最低为2.43 mmol m~(-2)s~(-1),且其结果枝光抑制参数最小,为0.629。说明开心形为最佳的高光效树形。(4)冠层微环境因子、叶片结构及光合生理指标之间多呈极显著相关关系,但开心形叶片结构和生理的大部分指标与冠层环境因子之间相关性较低,说明开心形树形不同部位的营养枝和结果枝的叶片性状差异较小,其光渗透性好,整个冠层的光截获能力和有效光辐射的分布差异较小,笔者认为开心形是龙安柚栽培中的适宜高光效树形。     

生产力调控对翌年苹果园土壤水分和苹果叶片光合特性的影响

以盛果期‘长枝富士’为试验材料, 以生产力调控翌年苹果(Malus pumila)树坐果期为主要观测期, 对不同生产力水平下果园土壤水分状况及苹果叶片光合特性进行了研究。结果表明: 在生产力调控范围内, 土壤剖面各个层次土壤含水量均随着生产力水平的减小而增加, 其中在60 cm处达到最大增量, 为31.02%; 而在600 cm范围内土壤贮水量最大能够提高15.41%。随着生产力水平的降低, “光合午休”现象减弱, 净光合速率(P_n)增加, 最大增幅为25.71%, 下午时段的蒸腾速率(T_r)下降迅速, 水分利用效率(WUE)最大提高34.12%。通过相关分析表明, 土壤贮水量(WSC)与P_n、T_r、WUE之间均达到显著相关, 其相关系数分别为: 0.973^**、-0.543^*和0.992^**。土壤贮水量(x)与水分利用效率(y)之间符合y = 0.002 3x - 1.480 6, R²= 0.984 4^**的回归模型。通过生产力调控可以改善土壤水分状况和果树光合能力, 提高WUE。     

不同光强对入侵种三裂叶豚草表型可塑性的影响

通过人工遮光,研究了不同光照强度下入侵植物三裂叶豚草的形态、生物量分配以及光合特性的表型可塑性.结果表明:与对照相比,遮光条件下,三裂叶豚草的株高、冠宽、单株叶面积、比叶面积和叶生物量比重显著增加,总生物量、单位叶面积生物量和根冠比减小;在全光照条件下,其冠宽和单株叶面积较小,根冠比较大,有利于高温强光下减少水分散失,表现出对不同光强较强的形态和生物量可塑性.遮光使三裂叶豚草叶片的日均净光合速率、蒸腾速率和气孔导度下降,胞间CO2浓度上升.正午光照最强时,低遮光处理植株的净光合速率、蒸腾速率和气孔导度最大.中遮光和高遮光条件下,其叶绿素含量显著增加,叶绿素a/b显著减小,有利于提高三裂叶豚草的光能利用效率,以适应弱光环境.     

紫耳箭竹克隆形态可塑性对典型冠层结构及光环境的响应

在重庆金佛山国家自然保护内,选择了3种典型群落类型(落叶阔叶林、常绿落叶阔叶混交林和常绿阔叶林),使用Hemiview数字植物冠层分析系统量化群落冠层结构和光环境特征,并对林下紫耳箭竹(Fargesia decurvata)的形态可塑性特征进行调查,分析冠层结构和光环境特征改变下紫耳箭竹形态可塑性的差异,并探讨它们之间的相互关系。结果表明:(1)随着落叶阔叶林"常绿落叶阔叶混交林"常绿阔叶林演替的进行,群落的冠层开度降低,叶面积指数增加,平均叶倾角变小,趋于水平化,冠层对光的截获能力提高,林下光照的强度降低(P0.05)。(2)随着光照强度的降低,紫耳箭竹分株矮小化,叶片变窄,生物量积累降低,但通过增大比茎长、叶面积率和比叶面积提高对光的利用效率,并增大分枝角度和比隔长有效适应弱光环境。(3)在光照条件差的常绿阔叶林下,紫耳箭竹降低对地下茎的投资,将较多的生物量用于秆的增高增长和叶片的生长;而在光照条件好的落叶阔叶林环境下,紫耳箭竹降低对枝、叶生物量的分配,则加大对地下茎的投资,可认为是克隆植物对水分资源所表现的一种觅食行为。研究表明,紫耳箭竹种群随着冠层结构的改变发生了明显的可塑性变化,这些可塑性变化是种群对冠层结构和光环境差异的适应性反应的结果,有利于增强种群对异质生境中光资源的获取和利用;群落内部可以通过调控冠层结构的改变协调和控制小径竹种群的发展。     

光照条件对‘寒富’苹果叶片结构和光合特性的影响

以‘寒富’苹果为试材,利用光学显微镜技术和气体交换等方法,研究了光照条件对盆栽和田间条件下‘寒富’苹果叶片结构和光合特性的影响.结果表明： 同种栽培方式下,与全光照条件相比,遮阴环境下‘寒富’苹果叶片的栅栏组织、海绵组织以及叶片总厚度均降低,其中,栅栏组织分别降低34.5%(盆栽)和25.0%(大田),叶片总厚度分别降低27.1%(盆栽)和18.3%(大田);遮阴环境下大田‘寒富’苹果叶片的光补偿点(LCP)最低,为(30.8±1.3） μmol·m^-2·s^-1,全光照比遮阴环境下叶片饱和光强分别高22.7%(盆栽)和48.2%(大田);盆栽不同光照环境下叶片对强光的适应能力不同,突然转入强光下,达到最大光合速率15.4 μmol·m^-2·s^-1(盆栽全光照)和12.7 μmol·m^-2·s^-1(盆栽遮阴)的光合启动时间不同,分别为23和33 min.表明长期遮阴影响‘寒富’苹果叶片质量及光合能力.     

不同海拔环境因子对富士苹果叶片和果实品质的影响

为了探究不同海拔生境富士苹果叶片和果实品质的差异及其对环境因子的响应,本文测定了3个海拔梯度(1375、1575、1715 m)上富士苹果叶片形态结构、解剖结构、δ13C等指标及果实品质,并应用逐步回归方法分析它们对环境因子的响应.结果表明:温暖指数、水热综合因子、光合有效辐射、寒冷指数、紫外线B和年降水量6个环境因子对叶片和果实品质特征参数起主导作用.随着海拔升高,温暖指数降低、水热综合因子增大、光合有效辐射增强、寒冷指数升高、紫外线B增强、年降水量增加,叶片结构和果实品质特征参数发生了不同程度的变化,具体表现为:叶片厚度、角质层厚度、栅海比、主脉最大导管直径、δ13C和单位面积叶片N含量逐渐增大,叶片长宽比、比叶面积、气孔长宽比和上下表皮占叶片厚度的比例逐渐减小;果形指数、果实硬度、糖酸比、色泽总量和色泽比逐渐增大,果实可滴定酸含量、色度角逐渐减小.随海拔升高,叶片光合速率增强,水分利用率增加,果实糖酸比呈上升趋势,高海拔比低海拔有相对较好的果实风味和外观着色,因此,在海拔1375~1715 m范围内,较高的海拔更有利于富士苹果生长.     

Modelling the effects of elevated atmospheric CO2 on crown development, light interception and photosynthesis of poplar in open top chambers

An open-top chamber experiment was carried out to examine the likely effects of elevated atmospheric [CO₂] on architectural as well as on physiological characteristics of two poplar clones ( Populus trichocarpa × P. deltoides clone Beaupré and P. deltoides × P. nigra clone Robusta). Crown architectural parameters required as input parameters for a three-dimensional (3D) model of poplar structure, such as branching frequency and position, branch angle, internode length and its distribution pattern, leaf size and orientation, were measured following growth in ambient and elevated [CO₂ ] (ambient + 350 μmol mol^–1) treated open-top chambers. Based on this information, the light interception and photosynthesis of poplar canopies in different [CO₂] treatments were simulated using the 3D poplar tree model and a 3D radiative transfer model at various stages of the growing season. The first year experiments and modelling results showed that the [CO₂] enrichment had effects on light intercepting canopy structure as well as on leaf photosynthesis properties. The elevated [CO₂] treatment resulted in an increase of leaf area, canopy photosynthetic rate and above-ground biomass production of the two poplar clones studied. However, the structural components responded less than the process components to the [CO₂] enrichment. Among the structural components, the increase of LAI contributed the most to the canopy light interception and canopy photosynthesis; the change of other structural aspects as a whole caused by the [CO₂] enrichment had little effect on daily canopy light interception and photosynthesis.     

Light interception in apple trees influenced by canopy architecture manipulation

Improvement of light penetration within tree canopies has been a constant objective of fruit tree architecture manipulation through the setting up of training systems. Recently, centrifugal training, i.e. the removal of fruiting shoots in the tree centre and on the underside of branches, has been proposed to improve fruit size and colour as well as return-bloom as compared to conventional solaxe-trained trees with equivalent crop loads. The present study was conducted to quantify the benefits of centrifugal training on light interception by the fruiting shoots via computer-assisted three-dimensional representations of foliage geometry. Data were collected on six 5-year-old apple trees cv.Galaxy, trained either with solaxe or centrifugal training systems, using an electromagnetic 3D digitiser. The 3D distribution of the foliage in the tree canopy was recreated by combining both the spatial locations of shoots (as measured from 3D digitising) and foliage reconstruction. Light interception efficiency properties of the trees were characterised by silhouette to total area ratio (STAR) values computed from images of the 3D mock-ups. Compared to the solaxe system, centrifugal training significantly improved the STAR of the whole tree by 20%. It also increased both leaf area and STAR of the fruiting shoots by approximately 15%, regardless of their position in the canopy. In this paper, we discuss the role of this enhanced light interception by the canopy in increasing the autonomy of the fruiting shoot, i.e. improved fruit size and colour, and return-bloom.     

Drought alters the canopy architecture and micro-climate of Hevea brasiliensis trees

 In this study a comparison of the canopy architecture and the growth and distribution of roots was made in 10-year-old trees of Hevea brasiliensis grown in a severely drought-prone area on the west coast of India under rainfed and irrigated conditions. LAI and light interception increased significantly in the irrigated compared to the rainfed trees. Girth and height of the tree were 29 and 19% more while width and height of the canopy were 19 and 20% more in the irrigated than rainfed trees. There were 22% more primary branches which had 26% more diameter in the irrigated trees than rainfed trees. The branches were inserted on the main trunk at an angle of 58.36° in the irrigated and 44.22° in rainfed trees. The above changes led to more light penetration which altered the light distribution inside the rainfed trees during summer and inhibited leaf photosynthesis particularly in the top canopy leaves. In the rainfed trees most of the growth occurred during the short favorable season immediately after the monsoon between June and October and no growth or even shrinking of the trunk was seen during summer. In the irrigated trees a higher growth was seen throughout the year and summer had no adverse effect. Although there was some difference in the root distribution pattern, the total root density per unit soil volume did not vary between the irrigated and rainfed trees. Key words  Hevea brasiliensis· Drought · Crown architecture · Micro-climate · Root growth Received: 8 May 1998 / Accepted 8 October 1998     

Optimal leaf area indices in C3 and C4 mono- and dicotyledonous species at low and high nitrogen availability

From an analytical model it was shown that for a given total amount of nitrogen in the canopy, there exists an optimal leaf area index (LAI), and therefore an optimal average leaf introgen content, at which canopy photosynthesis is maximal. If the LAI is increased above this optimum, increased light interception will not compensate for reduction in photosynthetic capacity of the canopy resulting from reduced leaf nitrogen contents. It was further derived from the model that the value of the optimal LAI increases with the photosynthetic nitrogen use efficiency (PNUE) and decreases with the canopy extinction coefficient for light (K_L) and incident photon flux density (PFD) at the top of the canopy. These hypotheses were tested on dense stands of species with different photosynthetic modes and different architectures. A garden experiment was carried out with the C₄ monocot sorghum ( Sorghum bicolor [L.] Moensch cv. Pioneer), the C₃ monocot rice ( Oryza sativa L. cv. Araure 4), the C₄ dicot amaranth ( Amaranthus cruentus L. cv. K113) and the C₃ dicot soybean ( Glycine max [L.] Merr. cv. Williams) at two levels of nitrogen availability.
The C₄ species had higher PNUEs than the C₃ species while the dicots formed stands with higher extinction coefficients for light and had lower PNUEs than the monocots. The C₄ and monocot species were found to have formed more leaf area per unit leaf nitrogen (i.e., had lower leaf nitrogen contents) than the C₃ and dicot species, respectively. These results indicate that the PNUE and the extinction coefficient for light are important factors determining the amount of leaf area produced per unit nitrogen as was predicted by the model.     

Simple equations to estimate light interception by isolated trees from canopy structure features: assessment with three-dimensional digitized apple trees

* Simple models of light interception are useful to identify the key structural parameters involved in light capture. We developed such models for isolated trees and tested them with virtual experiments. Light interception was decomposed into the projection of the crown envelope and the crown porosity. The latter was related to tree structure parameters. * Virtual experiments were conducted with three-dimensional (3-D) digitized apple trees grown in Lebanon and Switzerland, with different cultivars and training. The digitized trees allowed actual values of canopy structure (total leaf area, crown volume, foliage inclination angle, variance of leaf area density) and light interception properties (projected leaf area, silhouette to total area ratio, porosity, dispersion parameters) to be computed, and relationships between structure and interception variables to be derived. * The projected envelope area was related to crown volume with a power function of exponent 2/3. Crown porosity was a negative exponential function of mean optical density, that is, the ratio between total leaf area and the projected envelope area. The leaf dispersion parameter was a negative linear function of the relative variance of leaf area density in the crown volume. * The resulting models were expressed as two single equations. After calibration, model outputs were very close to values computed from the 3-D digitized databases.     

Exploring the spatial distribution of light interception and photosynthesis of canopies by means of a functional-structural plant model

Background and Aims

At present most process-based models and the majority of three-dimensional models include simplifications of plant architecture that can compromise the accuracy of light interception simulations and, accordingly, canopy photosynthesis. The aim of this paper is to analyse canopy heterogeneity of an explicitly described tomato canopy in relation to temporal dynamics of horizontal and vertical light distribution and photosynthesis under direct- and diffuse-light conditions.

Methods

Detailed measurements of canopy architecture, light interception and leaf photosynthesis were carried out on a tomato crop. These data were used for the development and calibration of a functional–structural tomato model. The model consisted of an architectural static virtual plant coupled with a nested radiosity model for light calculations and a leaf photosynthesis module. Different scenarios of horizontal and vertical distribution of light interception, incident light and photosynthesis were investigated under diffuse and direct light conditions.

Key Results

Simulated light interception showed a good correspondence to the measured values. Explicitly described leaf angles resulted in higher light interception in the middle of the plant canopy compared with fixed and ellipsoidal leaf-angle distribution models, although the total light interception remained the same. The fraction of light intercepted at a north–south orientation of rows differed from east–west orientation by 10 % on winter and 23 % on summer days. The horizontal distribution of photosynthesis differed significantly between the top, middle and lower canopy layer. Taking into account the vertical variation of leaf photosynthetic parameters in the canopy, led to approx. 8 % increase on simulated canopy photosynthesis.

Conclusions

Leaf angles of heterogeneous canopies should be explicitly described as they have a big impact both on light distribution and photosynthesis. Especially, the vertical variation of photosynthesis in canopy is such that the experimental approach of photosynthesis measurements for model parameterization should be revised.     

Carbon gain in a multispecies canopy: the role of specific leaf area and photosynthetic nitrogen-use efficiency in the tragedy of the commons

Models have been formulated for monospecific stands in which canopy photosynthesis is determined by the vertical distribution of leaf area, nitrogen and light. In such stands, resident plants can maximize canopy photosynthesis by distributing their nitrogen parallel to the light gradient, with high contents per unit leaf area at the top of the vegetation and low contents at the bottom. Using principles from game theory, we expanded these models by introducing a second species into the vegetation, with the same vertical distribution of biomass and nitrogen as the resident plants but with the ability to adjust its specific leaf area (SLA, leaf area:leaf mass). The rule of the game is that invaders replace the resident plants if they have a higher plant carbon gain than those of the resident plants. We showed that such invaders induce major changes in the vegetation. By increasing their SLA, invading plants could increase their light interception as well as their photosynthetic nitrogen-use efficiency (PNUE, the rate of photosynthesis per unit organic nitrogen). By comparison with stands in which canopy photosynthesis is maximized, those invaded by species of high SLA have the following characteristics: (1) the leaf area index is higher; (2) the vertical distribution of nitrogen is skewed less; (3) as a result of the supra-optimal leaf area index and the more uniform distribution of nitrogen, total canopy photosynthesis is lower. Thus, in dense canopies we face a classical tragedy of the commons: plants that have a strategy to maximize canopy carbon gain cannot compete with those that maximize their own carbon gain. However, because of this strategy, individual as well as total canopy carbon gain are eventually lower. We showed that it is an evolutionarily stable strategy to increase SLA up to the point where the PNUE of each leaf is maximized.     

Effects of defruiting on source-sink relationship, carbon budget, leaf carbohydrate content and water use efficiency of apple trees

Three-year-old apple ( Malus domestica Borkh. cv. Golden Delicious) trees with ca 5 m² leaf area each were grown in situ in lysimeters and defruited either in August. September or October. Net photosynthesis, dark respiration and water use efficiency were measured for 5 days prior to and after fruit removal. using tree canopy chambers and infrared gas analysis with a dedicated interface and software. Net photosynthesis of the tree canopy declined after defruiting. irrespective of the month of defruiting. The decline after fruit removal was 30-40% in August or September and 57% in October. After defruiting in August or September, trees respired 50–60% slower in the dark, but 39% faster after defruiting in October. The excessive respiratory losses after fruit removal in October. when the tree lost more carbon than it assimilated. may have been induced by translocation of carbohydrates from the leaves to the perennial woody parts of the tree and by the onset of leaf senescence. Fruit removal in August or September increased water use efficiency by 31–41%. but decreased water use efficiency in October by 75%     

苹果园光能截获率的数学模型

应用气象学原理推导了不同纬度、不同栽植行向、不同树形的苹果园光能截获率的数学模型,给出在保证树冠基部外围日照时间大于25％总日照时间的条件下,生长季中充分截获光能的最佳树形、行距、冠高、冠径的优化组合方案。实例分析表明,计算结果与专家经验基本一致。本研究为果园合理栽植,充分利用光能,合理整形修剪提供理论依据与参考方案。