Simulation of photon transport in a three-dimensional leaf: implications for photosynthesis

A model to evaluate photon transport within leaves and the implications for photosynthesis are investigated. A ray tracing model, Raytran, was used to produce absorption profiles within a virtual dorsiventral plant leaf oriented in two positions (horizontal/vertical) and illuminated on one of its two faces (adaxial/abaxial). Together with chlorophyll profiles, these absorption profiles feed a simple photosynthesis model that calculates the gross photosynthetic rate as a function of the incident irradiance. The differences observed between the four conditions are consistent with the literature: horizontal‐adaxial leaves, which are commonly found in natural conditions, have the greatest light use efficiency. The absorption profile obtained with horizontal‐abaxial leaves lies below this, but above those obtained for vertical leaves. The latter present similar gross photosynthetic rates when irradiated on either the adaxial or abaxial surfaces. Vertical profiles of photosynthetic rates across the leaf confirm that carbon fixation occurs mainly in the palisade parenchyma, that the leaf anatomy is integral to its function and that leaves cannot be considered as a single homogeneous unit. Finally, the relationships between leaf structure, orientation and photosynthesis are discussed.     

Profiles of light absorption and chlorophyll within spinach leaves from chlorophyll fluorescence

Chlorophyll fluorescence was used to estimate profiles of absorbed light within chlorophyll solutions and leaves. For chlorophyll solutions, the intensity of the emitted fluorescence declined in a log–linear manner with the distance from the irradiated surface as predicted by Beer's law. The amount of fluorescence was proportional to chlorophyll concentration for chlorophyll solutions given epi‐illumination on a microscope slide. These relationships appeared to hold for more optically complex spinach leaves. The profile of chlorophyll fluorescence emitted by leaf cross sections given epi‐illumination corresponded to chlorophyll content measured in extracts of leaf paradermal sections. Thus epifluorescence was used to estimate relative chlorophyll content through leaf tissues. Fluorescence profiles across leaves depended on wavelength and orientation, reaching a peak at 50–70 µm depth. By infiltrating leaves with water, the pathlengthening due to scattering at the airspace : cell wall interfaces was calculated. Surprisingly, the palisade and spongy mesophyll had similar values for pathlengthening with the value being greatest for green light (550 > 650 > 450 nm). By combining fluorescence profiles with chlorophyll distribution across the leaf, the profile of the apparent extinction coefficient was calculated. The light profiles within spinach leaves could be well approximated by an apparent extinction coefficient and the Beer–Lambert/Bouguer laws. Light was absorbed at greater depths than predicted from fibre optic measurements, with 50% of blue and green light reaching 125 and 240 µm deep, respectively.     

Ontogenetic differences in mesophyll structure and chlorophyll distribution in Eucalyptus globulus ssp. globulus

Mesophyll structure has been associated with the photosynthetic performance of leaves via the regulation of internal light and CO(2) profiles. Differences in mesophyll structure and chlorophyll distribution within three ontogenetically different leaf types of Eucalyptus globulus ssp. globulus were investigated. Juvenile leaves are blue-grey in color, dorsiventral (adaxial palisade layer only), hypostomatous, and approximately horizontal in orientation. In contrast, adult leaves are dark green in color, isobilateral (adaxial and abaxial palisade), amphistomatous, and nearly vertical in orientation. The transitional leaf type has structural features that appear intermediate between the juvenile and adult leaves. The ratio of mesophyll cell surface area per unit leaf surface area (A(mes)/A) of juvenile leaves was maximum at the base of a single, adaxial palisade layer and declined through the spongy mesophyll. Chlorophyll a + b content showed a coincident pattern, while the chlorophyll a:b ratio declined linearly from the adaxial to abaxial epidermis. In comparison, the mesophyll of adult leaves had a bimodal distribution of A(mes)/A, with maxima occurring beneath both the adaxial and abaxial surfaces within the first layer of multiple palisade layers. The distribution of chlorophyll a + b content had a similar pattern, although the maximum ratio of chlorophyll a:b occurred immediately beneath the adaxial and abaxial epidermis. The matching distributions of A(mes)/A and chlorophyll provide further evidence that mesophyll structure may act to influence photosynthetic performance. These changes in internal leaf structure at different life stages of E. globulus may be an adaptation for increased xeromorphy under increasing light exposure experienced from the seedling to adult tree, similar to the characteristics reported for different species according to sunlight exposure and water availability within their native habitats.     

Measurement of gradients of absorbed light in spinach leaves from chlorophyll fluorescence profiles

Profiles of chlorophyll fluorescence were measured in spinach leaves irradiated with monochromatic light. The characteristics of the profiles within the mesophyll were determined by the optical properties of the leaf tissue and the spectral quality of the actinic light. When leaves were infiltrated with 10^?4M DCMU [3‐(3,4‐dichlorophenyl)‐1, 1‐dimethyl‐urea] or water, treatments that minimized light scattering, irradiation with 2000 μmol m^?2 s^?1 green light produced broad Gaussian‐shaped fluorescence profiles that spanned most of the mesophyll. Profiles for chlorophyll fluorescence in the red (680 ± 16 nm) and far red (λ > 710 nm) were similar except that there was elevated red fluorescence near the adaxial leaf surface relative to far red fluorescence. Fluorescence profiles were narrower in non‐infiltrated leaf samples where light scattering increased the light gradient. The fluorescence profile was broader when the leaf was irradiated on its adaxial versus abaxial surface due to the contrasting optical properties of the palisade and spongy mesophyll. Irradiation with blue, red and green monochromatic light produced profiles that peaked 50, 100 and 150 μm, respectively, beneath the irradiated surface. These results are consistent with previous measurements of the light gradient in spinach and they agree qualitatively with measurements of carbon fixation under monochromatic blue, red and green light. These results suggest that chlorophyll fluorescence profiles may be used to estimate the distribution of quanta that are absorbed within the leaf for photosynthesis.     

Photosynthesis and resource distribution through plant canopies

Plant canopies are characterized by dramatic gradients of light between canopy top and bottom, and interactions between light, temperature and water vapour deficits. This review summarizes current knowledge of potentials and limitations of acclimation of foliage photosynthetic capacity (A(max)) and light-harvesting efficiency to complex environmental gradients within the canopies. Acclimation of A(max) to high light availability involves accumulation of rate-limiting photosynthetic proteins per unit leaf area as the result of increases in leaf thickness in broad-leaved species and volume: total area ratio and mesophyll thickness in species with complex geometry of leaf cross-section. Enhancement of light-harvesting efficiency in low light occurs through increased chlorophyll production per unit dry mass, greater leaf area per unit dry mass investment in leaves and shoot architectural modifications that improve leaf exposure and reduce within-shoot shading. All these acclimation responses vary among species, resulting in species-specific use efficiencies of low and high light. In fast-growing canopies and in evergreen species, where foliage developed and acclimated to a certain light environment becomes shaded by newly developing foliage, leaf senescence, age-dependent changes in cell wall characteristics and limited foliage re-acclimation capacity can constrain adjustment of older leaves to modified light availabilities. The review further demonstrates that leaves in different canopy positions respond differently to dynamic fluctuations in light availability and to multiple environmental stresses. Foliage acclimated to high irradiance respond more plastically to rapid changes in leaf light environment, and is more resistant to co-occurring heat and water stress. However, in higher light, co-occurring stresses can more strongly curb the efficiency of foliage photosynthetic machinery through reductions in internal diffusion conductance to CO(2). This review demonstrates strong foliage potential for acclimation to within-canopy environmental gradients, but also highlights complex constraints on acclimation and foliage functioning resulting from light x foliage age interactions, multiple environmental stresses, dynamic light fluctuations and species-specific leaf and shoot structural constraints.     

Photosynthetic stimulation under long-term CO2 enrichment and fertilization is sustained across a closed Populus canopy profile (EUROFACE)

The long-term response of leaf photosynthesis to rising CO2 concentrations [CO2] depends on biochemical and morphological feedbacks. Additionally, responses to elevated [CO2] might depend on the nutrient availability and the light environment, affecting the net carbon uptake of a forest stand. After 6 yr of exposure to free-air CO2 enrichment (EUROFACE) during two rotation cycles (with fertilization during the second cycle), profiles of light, leaf characteristics and photosynthetic parameters were measured in the closed canopy of a poplar (Populus) short-rotation coppice. Net photosynthetic rate (A(growth)) was 49% higher in poplars grown in elevated [CO2], independently of the canopy position. Jmax significantly increased (15%), whereas leaf carboxylation capacity (Vcmax), leaf nitrogen (N(a)) and chlorophyll (Chl(a)) were unaffected in elevated [CO2]. Leaf mass per unit area (LMA) increased in the upper canopy. Fertilization created more leaves in the top of the crown. These results suggest that the photosynthetic stimulation by elevated [CO2] in a closed-canopy poplar coppice might be sustained in the long term. The absence of any down-regulation, given a sufficient sink capacity and nutrient availability, provides more carbon for growth and storage in this bioenergy plantation.     

A new analytical model for whole-leaf potential electron transport rate

A new analytical model for the response of whole‐leaf potential electron transport rate (J) to light is presented. The model treats incident irradiance at the upper and lower leaf surfaces independently, describes transdermal profiles of light absorption and electron transport capacity explicitly, and calculates J by integrating the minimum of light‐ and capacity‐limited rates among paradermal chlorophyll layers. The capacity profile is assumed to be a weighted average of two opposed exponential profiles, each of which corresponds to the profile of light‐limited rate when only one surface is illuminated; the weights may take on any values, provided they sum to unity, so the model can describe leaves with a wide range of ‘preferred’ illumination regimes. By treating irradiance at either surface independently and assuming the capacity profile is fixed on short time scales, the model predicts observed effects of leaf inversion on light‐response curves and their apparent convexity. By assuming the capacity profile can adapt on developmental time scales, the model can predict the observed dependence of inversion effects on the growth lighting regime. It is suggested that the new model, which is mathematically compact and formally similar to the standard non‐rectangular hyperbola model for J, be used in place of the standard model in studies in which the effects of leaf angle or diffuse light fraction on gas exchange are of interest.     

Diaheliotropic leaf movement enhances leaf photosynthetic capacity and photosynthetic light and nitrogen use efficiency via optimising nitrogen partitioning among photosynthetic components in cotton (Gossypium hirsutum L.)

• Phototropic leaf movement of plants is an effective mechanism for adapting to light conditions. Light is the major driver of plant photosynthesis. Leaf N is also an important limiting factor on leaf photosynthetic potential. Cotton (Gossypium hirsutum L.) exhibits diaheliotropic leaf movement. Here, we compared the long‐term photosynthetic acclimation of fixed leaves (restrained) and free leaves (allowed free movement) in cotton.
• The fixed leaves and free leaves were used for determination of PAR, leaf chlorophyll concentration, leaf N content and leaf gas exchange. The measurements were conducted under clear sky conditions at 0, 7, 15 and 30 days after treatment (DAT).
• The results showed that leaf N allocation and partitioning among different components of the photosynthetic apparatus were significantly affected by diaheliotropic leaf movement. Diaheliotropic leaf movement significantly increased light interception per unit leaf area, which in turn affected leaf mass per area (LMA), leaf N content (N_A) and leaf N allocation to photosynthesis (N_P). In addition, cotton leaves optimised leaf N allocation to the photosynthetic apparatus by adjusting leaf mass per area and N_A in response to optimal light interception.
• In the presence of diaheliotropic leaf movement, cotton leaves optimised their structural tissue and photosynthetic characteristics, such as LMA, N_A and leaf N allocation to photosynthesis, so that leaf photosynthetic capacity was maximised to improve the photosynthetic use efficiency of light and N under high light conditions.

     

The impact of light intensity on shade-induced leaf senescence

Plants often have to cope with altered light conditions, which in leaves induce various physiological responses ranging from photosynthetic acclimation to leaf senescence. However, our knowledge of the regulatory pathways by which shade and darkness induce leaf senescence remains incomplete. To determine to what extent reduced light intensities regulate the induction of leaf senescence, we performed a functional comparison between Arabidopsis leaves subjected to a range of shading treatments. Individually covered leaves, which remained attached to the plant, were compared with respect to chlorophyll, protein, histology, expression of senescence-associated genes, capacity for photosynthesis and respiration, and light compensation point (LCP). Mild shading induced photosynthetic acclimation and resource partitioning, which, together with a decreased respiration, lowered the LCP. Leaf senescence was induced only under strong shade, coinciding with a negative carbon balance and independent of the red/far-red ratio. Interestingly, while senescence was significantly delayed at very low light compared with darkness, phytochrome A mutant plants showed enhanced chlorophyll degradation under all shading treatments except complete darkness. Taken together, our results suggest that the induction of leaf senescence during shading depends on the efficiency of carbon fixation, which in turn appears to be modulated via light receptors such as phytochrome A.     

光合作用对光和二氧化碳响应的观测方法探讨

用便携式光合仪LI-6400观测自然条件下生长的盆栽蚕豆叶片光合作用对光和二氧化碳的响应发现：（1）用未经过光合诱导的叶片观测光合作用对光的响应会得到即使在全太阳光强下光合作用仍然不饱和的假象;（2）利用某些经验方程计算的饱和光强远低于实际观测值;（3）在观测光合作用对CO2的响应过程中,每一次CO2浓度变化都应当伴随一次光合仪的匹配步骤,否则所得结果偏差很大;（4）在不饱和光下观测光合作用对CO2的响应,会导致对叶片光合能力的低估。     

Why abaxial illumination limits photosynthetic carbon fixation in spinach leaves

Limitations of carbon fixation within spinach leaves due to light and CO2 were investigated. Under equivalent photon fluxes, carbon fixation was higher when leaves were irradiated adaxially compared to abaxially. Maximal carbon fixation occurred in the middle of the palisade mesophyll under adaxial illumination, whereas, maximal carbon fixation occurred in the spongy mesophyll under abaxial illumination. Total carbon fixation and the pattern of carbon fixation across leaves were similar, when leaves were irradiated with 800 micromol quanta m(-2) s(-1) either adaxially alone or adaxially plus abaxially (1,600 micromol quanta m(-2) s(-1)). In contrast, when both leaf surfaces were irradiated simultaneously with 200 micromol quanta m(-2) s(-1), total carbon fixation increased and carbon fixation in the middle of the leaf was higher compared to leaves irradiated unilaterally with the low light. Feeding 14CO2 through either the adaxial or abaxial leaf surface did not change the pattern of carbon fixation across the leaf. Increasing 14CO2 pulse-feeding times from 5 s to 60 s allowed more 14CO2 to be fixed but did not change the pattern of 14CO2 fixation across the leaf. We concluded that in spinach, the distribution of both light and Rubisco activity within leaves has significant effects on the patterns of carbon fixation across leaves; whereas CO2 diffusion does not appear to affect the carbon fixation pattern within spinach leaves.     

Profiles of photosynthetic oxygen-evolution within leaves of Spinacia oleracea

Oxygen evolution was measured from mesophyll tissues in spinach leaves using a photoacoustic technique. The photosynthetic capacity of individual cell layers was measured by directing microscopic beams of light, 40 μm wide, to cells exposed within a leaf cross section. The resulting profile for oxygen-evolution potential was relatively flat, indicating a uniform capacity for photosynthesis in leaf mesophyll tissues. Two experimental approaches were used to estimate the photosynthetic performance of individual mesophyll cell layers when white light was applied to the adaxial leaf surface. These experiments indicated that oxygen was produced relatively uniformly across the mesophyll and that oxygen evolution increased with irradiance of the white light applied to the leaf surface. The measured profiles for oxygen evolution and capacity are flatter than previous measurements of profiles of fixed carbon and estimates of profiles for absorbed light within spinach leaves.     

枫香叶片衰老过程中光合能力的变化

利用Li-6400XT便携式光合作用测定系统,于2014年10—12月测定枫香叶片衰老过程中光合作用光响应曲线,采用叶氏模型和非直角双曲线模型进行模拟,分析枫香叶片衰老过程中光合能力的变化.结果表明: 随着枫香叶片逐渐变黄变红,其净光合速率的光响应能力逐渐降低,实测的最大净光合速率从叶片开始泛黄时的2.88 μmol CO₂·m^-2·s^-1下降到叶片衰老后期(12月8日)的0.95 μmol CO₂·m^-2·s^-1.2种光响应模型均较好地模拟了观测的光响应数据,其中叶氏模型表现更优.模拟得到的最大净光合速率、表观量子效率、光补偿点的量子效率、暗呼吸速率等参数均随枫香叶片衰老凋落而逐渐下降,反映出枫香叶片衰老过程中光合能力缓慢下降的过程.在树梢红叶未落期间,枫香叶片仍具有一定的净光合作用能力,这有利于增加秋冬季节的碳吸收量.     

The flexible interrelation between AOX respiratory pathway and photosynthesis in rice leaves.

Alternative respiratory pathway was investigated in rice seedlings grown under total darkness, light/dark cycle, or continuous light. The capacity of the alternative pathway was relatively higher in leaves that had longer light exposure. An analysis of rice AOX1 multigene family revealed that AOX1c, but not AOX1a and AOX1b, had a light-independent expression. The alternative oxidase (AOX) inhibitor, salicylhydroxamic acid (SHAM, 1mM), inhibited nearly 68% of the capacity of the alternative pathway in leaves grown under different light conditions. The plants grown under different light periods were treated with SHAM and then were exposed to illumination for 4h. The transition from dark to 4h of light stimulated the capacity of alternative pathway in etiolated rice seedlings and in those grown under light/dark cycle, whereas the capacity of the alternative pathway was constant in seedlings grown under continuous light with additional 4h of illumination. Etiolated leaves did not show any CO(2) fixation after 4h of illumination, and the increase in chlorophyll content was delayed by the SHAM pretreatment. When seedlings grown under light/dark cycle were moved from dark and exposed to 4h of light, increases in chlorophyll content and CO(2) fixation rate were reduced by SHAM. Although these parameters were stable in plants grown under continuous light, SHAM decreased CO(2) fixation rate but not the chlorophyll content. These results indicate that the role and regulation of AOX in light are determined by the developmental stage of plant photosynthetic apparatus.     

普通野生稻和武育粳8号剑叶光合功能衰退的比较

以武育粳 8号 (Wuyujing8)和普通野生稻 (Oryza.sativaL .f.spontaneaRoschev .)为材料 ,比较了它们剑叶光合功能衰退的进程。结果表明 ,在剑叶一生中 ,武育粳 8号剑叶的叶绿素含量和光合速率都高于普通野生稻 ,光合速率的最高值前者为 2 3 .5 μmolCO2 m 2 ·S 1,后者为 1 5 .5 μmolCO2 m 2 ·S 1。前者剑叶光合速率的高值持续期比后者长近 2 0d ,其叶源量是后者的 2 .8倍。武育粳 8号剑叶叶绿素含量最大值的SPAD值为 5 0 .1 ,而普通野生稻最大值的SPAD值为 42 .7。在可溶性蛋白质的基础上 ,核酮糖 1 ,5 二磷酸羧化酶比活性二者之间没有太大区别。内肽酶活力上升落后于光合功能的衰退。在光合功能衰退的早期 ,内肽酶的活力仅有小幅度提高 ,但在叶片衰老的末期升高幅度则较大。这说明内肽酶主要在光合功能衰退的后期即不可逆衰退期才发挥较大作用。     

遮荫对甜瓜叶片光合特性的影响

以适宜不同栽培条件的3个甜瓜品种为材料,研究了遮荫对其光合色素含量、净光合速率及比叶重等叶片特性的影响。结果表明:与正常光照相比,遮光处理能诱使甜瓜叶片叶绿素和类胡萝卜素含量显著提高,叶绿素a/b显著降低。全光照条件下,3个甜瓜品种的净光合速率日变化曲线均呈双峰型,有明显的"光合午休"现象,而在遮荫条件下则呈单峰曲线变化,且光合峰值出现的时间比全光照下推迟;遮光条件下甜瓜叶片气孔导度日变化曲线与其净光合速率日变化类似。3个品种间比叶重在全光照条件下差异不显著,但遮荫显著降低了壮龄叶片的比叶重,且遮荫强度越重,比叶重越小;品种‘黄河蜜3号’壮龄叶的比叶重降幅(31.83%)显著大于‘银帝’(27.22%)和‘玉金香’(26.01%)。可见,遮荫降低了甜瓜叶片的净光合速率和功能叶片的比叶重,植株通过增加自身叶片光合色素含量以增强对环境的适应性,缓解遮荫对其的影响,品种‘银帝’表现出较强的耐弱光性。     

Canopy structure,vertical radiation profile and photosynthetic function in a Quercus ilex evergreen forest

The studied evergreen forest dominated by Quercus ilex showed a leaf area index (LAI) of 4.5, of which 61 % was accumulated within the tree layer, 30 % within the shrub layer, and 9 % within the herb layer. The leaves of all the species were ± horizontally oriented (41°), absorbing a relevant percentage of incident irradiance. The high LAI drastically modified the quality and quantity of solar radiation on the forest underground. The spectral distribution of the radiation under the forest was markedly deficient in blue and red wavelengths. The maximum absorption in these spectral bands was found in spring, when net photosynthetic rate (P _N ) was at its maximum, and in summer, when new leaves reached 90 % of their definitive structure. The vertical radiation profile showed an evident reduction of the red-far red ratio (R/FR). Radiation quality and quantity influenced leaf physiology and morphology. Clear differences in leaf size, leaf water content per area (LWC) and specific leaf area (SLA) on the vertical profile of the forest were observed. All the shrub species showed similar SLA (12.02 m2 kg-1, mean value). The ability to increase SLA whilst simultaneously reducing leaf thickness maximized the carbon economy. The high chlorophyll (Chl) content of shrub layer leaves (1.41 g kg-1, mean value) was an expression of shade adaptation. Both leaf morphology and leaf physiology expressed the phenotypic plasticity. Q. ilex, Phillyrea latifolia and Pistacia lentiscus of the forest shrub layer showed wide differences in leaf structure and function with respect to the same species developing under strong irradiance (low maquis): a 57 % mean increase of SLA and a 86 % mean decrease of PN. They showed high leaf plasticity. Leaf plasticity implies that the considered sclerophyllous species has an optimum developmental pattern achieving adaptation to environments. This revised version was published online in September 2006 with corrections to the Cover Date.     

The consequences of delayed greening during leaf development for light absorption and light use efficiency

Many species of rainforest plants have an unusual form of leaf development such that leaves delay greening until after full leaf expansion. Chlorophyll accumulation was measured during leaf development in five woody rainforest species, three with white young leaves, and two with ‘normal’ greening. In the three species with white leaves, the chlorophyll content of the expanding leaves was about 0.4mg dm^?2, whereas in the two species with green young leaves, chlorophyll content was about 2.1 mg dm^?2. Chlorophyll accumulation in greenhouse and field experiments was independent of light level. During leaf expansion, species with delayed chloroplast development only absorb 18–25% of the maximum possible light, compared with 80% for species with normal greening. Furthermore, species with delayed greening have low chlorophyll contents and reduced absorption for at least 30 d after full expansion. At a PPFD typical of the forest under story, the photosynthetic light use efficiency based upon incident radiation was 0.030–0.036 for species with delayed chloroplast development and 0.068–0.085 for the two species with normal greening. The lower light use efficiency of white species was primarily due to decreased light absorption. However, they also had a slightly lower light use efficiency based upon absorbed radiation, suggesting that development of other components of the photo-synthetic apparatus also may be delayed. Despite the fact that delayed greening decreases light absorption and light use efficiency during leaf development, it is extremely common in shade-tolerant species. We suggest that an advantage of delayed greening is that resources are not invested in the leaf until it is fully expanded and better defended from herbivores.     

A system for imaging transverse distribution of scattered light and chlorophyll fluorescence in intact rice leaves

In order to examine the transverse distribution of scattered light and chlorophyll fluorescence in intact rice leaves, a micro-fluorescence imaging system was devised using a microscope, a CCD camera with an image intensifier, an Ar and a He-Ne laser light source, an image processor, and a microcomputer. A laser light was projected vertically on to the surface of a rice leaf segment at a cut-edge, and scattered light and induced fluorescence were observed at the cut-section from a 90° angle to the axis of the laser beam. The intensity of scattered light showed a maximum at several micrometres depth from the leaf surface and a steep gradient afterwards. Fluorescence reached a maximum crossing with the decline curve of the scattered light. The maximum of fluorescence measured at 741 nm was observed at a greater depth from the leaf surface than that at 687 nm, suggesting that part of the fluorescence of the longer wavelength was emitted due to absorption of fluorescence of the shorter wavelength. Profiles of the scattered light and the chlorophyll fluorescence depended on leaf anatomy.