定向种植对夏玉米群体内光环境及叶片光合性能的影响

为改善玉米群体内光环境,进一步提高玉米单株光合能力以获得高产,本研究以郑单958为试验材料,通过设置种子定向入土方式,研究了定向有序种植条件下群体内光分布特征,以及单株玉米穗位叶花后光合性能,并借助快速叶绿素荧光动力学曲线分析了与叶片光合性能有密切联系的光系统Ⅱ(PSⅡ)的性能特征.结果表明:叶片不同朝向显著改变夏玉米群体内穗位叶处光合有效辐射截获量,朝南处理(S)平均比朝北处理(N)高271.8%.不同朝向的叶片对高光与弱光的利用能力差异显著,朝南处理饱和光强下净光合速率(Pn)显著升高,表明其高光强利用能力显著提升;而朝北处理(N)表观量子效率(α)则随生育期推进显著增加,有利于叶片适应长期弱光环境.生育前期朝南处理PSⅡ电子供体侧和受体侧性能显著提高,进而改善了PSⅡ反应中心性能(PIABS)和荧光光化学淬灭系数(ψo),电子在电子传递链中转移效率(φEo)的提高增强了电子由PSⅡ向光系统Ⅰ(PSⅠ)的传递性能.生育前期叶片性能呈现出朝南>朝东>朝西>朝北的趋势.但成熟末期朝南处理对强光的利用效率显著降低,朝北处理在生育后期表现出较强的弱光利用能力,表观量子效率显著升高,花后40dPn与PSⅡ性能均表现为朝北>朝西>朝东>朝南的趋势.总体上,朝南与朝东处理群体内光环境改善显著,群体穗位层截获光合有效辐射较多,光合能力和干物质生产能力增强,有利于夏玉米产量提高.     

绒毛番龙眼对生长光强的形态和生理适应

在100％、50％、25％和8％自然光强下栽培绒毛番龙眼幼苗并研究了其对光环境的适应。100％生长光强下绒毛番龙眼通过增大叶片悬挂角(midrib angle,MA)和比叶重(lamina mass per unit area,LMA),减少叶氮在捕光组分中的分配等降低光能捕获;通过增加类胡萝卜素含量增加热耗散。虽如此,还是发生了比较严重的光抑制,加之叶氮在光合机构中的分配最少,导致光合能力最低,长势最差。8％生长光强下绒毛番龙眼通过降低MA、LMA以及叶片技转,增加叶氮在捕光组分中的分配等提高光能捕获能力,光能转换及利用效率较高,热耗散水平较低,但由于环境光较弱,限制了光合碳同化,植株生长也较慢。50％和25％生长光强下绒毛番龙眼有较强的光能捕获、利用和耗散能力,在几种光处理中长势最好。     

新疆超高产棉花冠层光分布特征及其与群体光合生产的关系

以新疆超高产棉田(皮棉产量在4000 kg·hm^-2以上)为研究对象,分析不同生育时期棉花冠层光分布、群体光合速率和干物质积累量的变化,研究不同产量水平棉田冠层的光环境变化特征及其与群体光合生产的关系.结果表明： 超高产田盛花期到盛铃后期冠层上、中、下层光吸收率的比例为2∶2∶1,呈均匀分布,群体散射辐射和直射辐射透过系数分别为0.20～0.55和0.22～0.56,处于较适宜范围,中、下层叶片受光良好,冠层各层次叶片群体光合速率差异较小.与高产(3500 kg·hm^-2)和一般高产(3000 kg·hm^-2)棉田相比,超高产田在盛铃前期具有较高的叶面积指数和群体光合速率峰值,在初絮期和盛絮期的叶面积指数下降缓慢,群体光合速率峰值仍保持较高值,非叶绿色器官对产量形成的光合贡献增大,群体干物质积累量较高.在栽培过程中,调节冠层结构,使垂直方向上光辐射和群体光合能力分布均匀是确保棉花高效利用光能、实现超高产的重要途径.     

叶片叶肉结构对环境光强的适应及对光合作用的影响

本文利用Kubelka-Munk理论描述了平行光在叶片内的吸收和散射,同时利用叶片分层光合作用非直角双曲线光反应模型,给出了整张叶片光合作用计算式。最后利用优化理论阐明了叶片叶肉分化成光合特性具有明显差异的栅栏组织和海绵组织可能是对叶片内光梯度的一种适应;同时证明了叶片叶肉在一定环境光强下存在一个最佳的栅栏组织和海绵组织比例,并且这个比例随环境光强增大而增大,这最佳比例也受叶肉组织光合特性差异的影响。     

中甸角蒿光合作用对生长光强的响应

在原生地和引种地对高山花卉中甸角蒿（1ncarvillea zhongdianensis）光合作用和叶片性状对生长光强的响应进行研究。结果表明：在香格里拉,光合速率（Pn）、类胡萝卜素（Car）、色素比（Chla/b）均随光强的降低而下降;而比叶面积（SLA）、叶绿素b（Chlb）、叶氮含量（LNCa）随光强的降低而上升。中甸角蒿主要是通过叶片形态、生化效率和叶片氮分配来响应生长光强的变化,对生长光强的适应表现出较大可塑性,使得其相对比较容易引种驯化。中甸角蒿在香格里拉对光照具有较广的适应幅,从香格里拉移栽到昆明后,虽然Pn下降约10%,但RGR增加约30%,表明其可以在昆明较好生长。     

光环境对胡桃楸幼苗生长与光合作用的影响

为了解胡桃楸幼苗对光的需求及适应规律,采用Li-6400便携式光合测定系统研究了不同光环境处理（100%、60%、30％和15%自然光）条件下3年生胡桃楸幼苗（适应1年后）叶片光合能力的季节变化及其对光强的响应.结果表明：在春季,胡桃楸幼苗对光反应不敏感,夏季和秋季随着光强的增加,叶片的最大光合速率、最大羧化速率和最大电子传递速率均显著增加（P<0.05）.光饱和点随光强的下降而降低（P<0.05）,表观量子效率、暗呼吸速率和光补偿点在不同光环境下未发现显著差异.100%和60%自然光处理的幼苗相对生长率差异不显著,但是随着光强下降,相对生长率显著下降（P<0.05）,为60%>30％>15%自然光处理.胡桃楸幼苗对不同的光环境表现出较强的适应性和可塑性;同时,通过降低光饱和点和减少碳积累,也能适应15%～30％自然光环境.     

水稻群体光合生产能力的强化及其调控

对进一步强化水稻群体光合生产能力及其调控进行了论述。认为,在保证较高适宜叶面积指数的基础上,进一步优化群体结构和选育具广幅光强适应能力的品种是实现进一步强化水稻群体光合生产能力的关键。同时,在栽培上做到合理密植、提高施肥水平、提高粒／叶比、培育强大根群和加强后期田间管理。     

荒漠植物梭梭群体和叶片水平气体交换对不同土壤水分的响应

为揭示C4荒漠植物梭梭的抗旱性和适应环境的光合作用特征,在人工控制土壤水分条件下,选择代表性植株,使用改进同化箱和LI-8100土壤碳通量自动测量系统组成的群体光合作用测量系统进行测量.该系统能够自动、连续观测,在测量时光、温环境因子稳定,能够准确测定植物的群体水平光合作用.使用LI-6400测定叶片水平光合作用.在相同土壤水分条件下,群体与叶片水平光合速率存在极显著差异（P〈0.001）,当土壤水分为田间持水量的50%（土壤含水量约10%）左右时梭梭的光合能力最强,群体光合速率（CAP）日均值为6.22μmolCO2m-2s-1,叶片光合速率（Pn）日均值为20.18μmolCO2m-2s-1,群体为叶片水平的30.8%.升高或降低土壤水分,梭梭的光合能力都下降.CAP与Pn的线性回归关系为CAP=0.20Pn＋1.82（r2=0.89,P〈0.001）.结果表明,适宜的土壤含水量可显著提高梭梭群体和叶片的光合能力,过高的土壤含水量不利于梭梭生长发育.梭梭群体及叶片水平的气体交换对同一水分条件有近似的响应趋势,利用拟合公式,可从叶片水平推算出群体水平的光合速率.     

大气CO2浓度和温度升高对水稻叶片及群体光合作用的影响

大气ＣＯ２浓度升高对植物光合作用的影响研究多集中在单叶水平,在高ＣＯ２及高温下对植物单叶及群体光合进行比较的研究少有报道,而群体水平的研究则是预测生态系统反应所不可缺少的。采用田间开顶式培养室研究了大气ＣＯ２浓度和温度升高对水稻（ＯｒｙｚａｓａｔｉｖａＬ．）叶片及群体光合作用的影响。发现ＣＯ２浓度和温度对水稻叶片光合作用有协同促进作用,而对群体光合作用的促进则随时间的推移而减弱;单叶光合受到的促进作用大于群体光合;叶面积指数只在营养生长期受到促进,冠层叶片含氮量受ＣＯ２影响降低。群体呼吸（包括茎杆）增加及冠层叶片早衰可能是后期ＣＯ２对群体光合促进作用下降的原因。     

水稻群体冠层光分布及光合作用模型

作物冠层光分布及光合作用模型,是作物栽培学、作物育种学研究的共同基础,对优化设计和评价作物株型、模拟作物生长发育与对环境变化的响应研究都有十分重要的价值.本文根据水稻群体冠层结构的特点,在虚拟切层法的基础上,建立了水稻群体冠层光分布及光合速率模型,模型包括冠层形态子模型、冠层光分布子模型和冠层光合速率子模型等.利用本模型,对设定的15625种水稻株型的光合速率进行了模拟计算,获得水稻最佳株型模型.结果表明,水稻群体光合速率与叶片数、叶含氮量、叶长、叶宽和叶倾角等5因素密切相关;最佳株型的上述5因素在冠层上部取值大,向下逐渐变小.     

Dynamics of Vertical Leaf Nitrogen Distribution in a Vegetative Wheat Canopy. Impact on Canopy Photosynthesis

The development of vertical canopy gradients of leaf N has beenregarded as an adaptation to the light gradient that helps tomaximize canopy photosynthesis. In this study we report thedynamics of vertical leaf N distribution during vegetative growthof wheat in response to changes in N availability and sowingdensity. The question of to what extent the observed verticalleaf N distribution maximized canopy photosynthesis was addressedwith a leaf layer model of canopy photosynthesis that integratesN-dependent leaf photosynthesis according to the canopy lightand leaf N distribution. Plants were grown hydroponically attwo amounts of N, supplied in proportion to calculated growthrates. Photosynthesis at light saturation correlated with leafN. The vertical leaf N distribution was associated with thegradient of absorbed light. The leaf N profile changed duringcrop development and was responsive to N availability. At highN supply, the leaf N profiles were constant during crop development.At low N supply, the leaf N profiles fluctuated between moreuniform and steep distributions. These changes were associatedwith reduced leaf area expansion and increasing N remobilizationfrom lower leaf layers. The distribution of leaf N with respectto the gradient of absorbed irradiance was close to the theoreticaloptimum maximizing canopy photosynthesis. Sensitivity analysisof the photosynthesis model suggested that plants maintain anoptimal vertical leaf N distribution by balancing the capacityfor photosynthesis at high and low light. Copyright 2000 Annalsof Botany Company Canopy photosynthesis, leaf nitrogen distribution, nitrogen, Triticum aestivum L, wheat     

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.     

A model of dynamics of leaves and nitrogen in a plant canopy: an integration of canopy photosynthesis,leaf life span,and nitrogen use efficiency

A model of dynamics of leaves and nitrogen is developed to predict the effect of environmental and ecophysiological factors on the structure and photosynthesis of a plant canopy. In the model, leaf area in the canopy increases by the production of new leaves, which is proportional to the canopy photosynthetic rate, with canopy nitrogen increasing with uptake of nitrogen from soil. Then the optimal leaf area index (LAI; leaf area per ground area) that maximizes canopy photosynthesis is calculated. If leaf area is produced in excess, old leaves are eliminated with their nitrogen as dead leaves. Consequently, a new canopy having an optimal LAI and an optimal amount of nitrogen is obtained. Repeating these processes gives canopy growth. The model provides predictions of optimal LAI, canopy photosynthetic rates, leaf life span, nitrogen use efficiency, and also the responses of these factors to changes in nitrogen and light availability. Canopies are predicted to have a larger LAI and a higher canopy photosynthetic rate at a steady state under higher nutrient and/or light availabilities. Effects of species characteristics, such as photosynthetic nitrogen use efficiency and leaf mass per area, are also evaluated. The model predicts many empirically observed patterns for ecophysiological traits across species.     

Optimal nitrogen distribution within a leaf canopy under direct and diffuse light

Nitrogen distribution within a leaf canopy is an important determinant of canopy carbon gain. Previous theoretical studies have predicted that canopy photosynthesis is maximized when the amount of photosynthetic nitrogen is proportionally allocated to the absorbed light. However, most of such studies used a simple Beer's law for light extinction to calculate optimal distribution, and it is not known whether this holds true when direct and diffuse light are considered together. Here, using an analytical solution and model simulations, optimal nitrogen distribution is shown to be very different between models using Beer's law and direct–diffuse light. The presented results demonstrate that optimal nitrogen distribution under direct–diffuse light is steeper than that under diffuse light only. The whole‐canopy carbon gain is considerably increased by optimizing nitrogen distribution compared with that in actual canopies in which nitrogen distribution is not optimized. This suggests that optimization of nitrogen distribution can be an effective target trait for improving plant productivity.     

Acclimation of photosynthesis in canopies: models and limitations

Within a time-scale of several days photosynthesis can acclimate to light by variation in the capacity for photosynthesis with depth in a canopy or by variation in the stoichiometry of photosynthetic components at each position within the canopy. The changes in leaf photosynthetic capacity are usually related to and expressed as changes in leaf nitrogen content. However, photosynthetic capacity and leaf nitrogen never match exactly the photon flux density (PFD) gradient within a canopy. As a result, photosynthetic light use efficiency, i.e. photosynthetic performance per incident PFD, increases considerably from the top of the canopy to the lower shaded part. Many of existing optimisation models fail to express the actual pattern of nitrogen or photosynthetic capacity distribution within a canopy. This failure occurs because these optimisation models do not consider that the quantitative aspect of photosynthesis acclimation is a whole plant phenomenon. Although turnover models, which describe the distribution of the photosynthetic apparatus within a canopy as a dynamic equilibrium between breakdown and regeneration of apparatus with respect to nitrogen availability, photosynthetic rate and export of carbohydrates, produce realistic results, these models require confirmation. The mechanism responsible for changes in the relative share of light-harvesting apparatus as acclimation to irradiance remains unknown. Ability of the photosynthetic apparatus to balance properly the light harvesting capacity with electron transport and biochemical capacities is limited. As a result of this fundamental limitation, photosynthetic light use efficiency always increases with increasing thickness of the photosynthetic apparatus.     

Maximizing daily canopy photosynthesis with respect to the leaf nitrogen allocation pattern in the canopy

Summary A model of daily canopy photosynthesis was constructed taking light and leaf nitrogen distribution in the canopy into consideration. It was applied to a canopy of Solidago altissima. Both irradiance and nitrogen concentration per unit leaf area decreased exponentially with increasing cumulative leaf area from the top of the canopy. The photosynthetic capacity of a single leaf was evaluated in relation to irradiance and nitrogen concentration. By integration, daily canopy photosynthesis was calculated for various canopy architectures and nitrogen allocation patterns. The optimal pattern of nitrogen distribution that maximizes the canopy photosynthesis was determined. Actual distribution of leaf nitrogen in the canopy was more uniform than the optimal one, but it realized over 20% more photosynthesis than that under uniform distribution and 4.7% less photosynthesis than that under the optimal distribution. Redeployment of leaf nitrogen to the top of the canopy with ageing should be more effective in increasing total canopy photosynthesis in a stand with a dense canopy than in a stand with an open canopy.     

Variation in crown light utilization characteristics among tropical canopy trees

BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.     

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

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

棉花叶片氮含量的空间分布与光合特性

在棉花生长旺季,将冠层按高度分多层测定了田间叶片含氮量和叶片净光合速率对光合有效辐射通量密度的响应(光响应曲线,Pn-PPFD response curve)及相应的生物指标.结果表明,各层叶片氮含量与光合作用关系密切,各层平均值大小依次为上层＞中层＞下层,对应层叶片的最大净光合速率Pmax、表观暗呼吸速率Rd、光补偿点LCP及光饱和点LSP均从上到下依次递减,与氮含量分布一致,而表观光合量子效率AQY则略有不同;氮含量的指数衰减系数 kn =0.762(R2=0.593),根据观测结果,棉田叶片氮含量(N)空间分布可以用相对累积叶面积指数(Lc/Lt)为自变量的指数方程来模拟,从而为建立光合作用机理模型与进行生产力奠定基础.     

Light and VPD gradients drive foliar nitrogen partitioning and photosynthesis in the canopy of European beech and silver fir

While foliar photosynthetic relationships with light, nitrogen, and water availability have been well described, environmental factors driving vertical gradients of foliar traits within forest canopies are still not well understood. We, therefore, examined how light availability and vapour pressure deficit (VPD) co-determine vertical gradients (between 12 and 42 m and in the understorey) of foliar photosynthetic capacity (Amax), 13C fractionation (∆), specific leaf area (SLA), chlorophyll (Chl), and nitrogen (N) concentrations in canopies of Fagus sylvatica and Abies alba growing in a mixed forest in Switzerland in spring and summer 2017. Both species showed lower Chl/N and lower SLA with higher light availability and VPD at the top canopy. Despite these biochemical and morphological acclimations, Amax during summer remained relatively constant and the photosynthetic N-use efficiency (PNUE) decreased with higher light availability for both species, suggesting suboptimal N allocation within the canopy. ∆ of both species were lower at the canopy top compared to the bottom, indicating high water-use efficiency (WUE). VPD gradients strongly co-determined the vertical distribution of Chl, N, and PNUE in F. sylvatica, suggesting stomatal limitation of photosynthesis in the top canopy, whereas these traits were only related to light availability in A. alba. Lower PNUE in F. sylvatica with higher WUE clearly indicated a trade-off in water vs. N use, limiting foliar acclimation to high light and VPD at the top canopy. Species-specific trade-offs in foliar acclimation to environmental canopy gradients may thus be considered for scaling photosynthesis from leaf to canopy to landscape levels.