Acclimation of photosynthesis to light and canopy nitrogen distribution: an interpretation

BACKGROUND AND AIMS: Acclimation of photosynthesis to light and its connection with canopy nitrogen (N) distribution are considered. An interpretation of a proportionality between light-saturated photosynthesis and local averaged leaf irradiance is proposed by means of a simple model. MODEL: The model assumes (a) local irradiance drives synthesis of photosynthetic protein from metabolic N; (b) photosynthetic N is slowly degraded over approx. 5-7 d; (c) metabolic N is equally available through the canopy. CONCLUSIONS: The kinetics of acclimation at different light levels may provide a way of parameterizing and testing the model. The model provides a rationale for the proportionality assumption mentioned above, which, while it is consistent with much experimental work, is valuable because it allows canopy photosynthesis to be calculated analytically.     

Expansins in growing tomato leaves

An expansin-like protein from growing tomato leaves was identified by its ability to restore the 'acid-growth' response to heat-inactivated tomato walls and by its similarity to expansins from cucumber hypocotyls. Native walls from growing tomato leaves exhibit an endogenous acid-induced extension (creep) that resembles in various biochemical characteristics the acid-growth activity of cucumber hypocotyls. For example, the acid-growth activity is lost when the walls of tomato leaves are briefly heated and is largely restored by addition of a crude protein extract from the walls of growing leaves. Wall proteins from growing leaves enhance the stress relaxation spectrum of tomato walls in a fashion characteristic of cucumber expansins. HPLC fractionation of the crude wall protein from tomato leaves yielded an active fraction containing a major 27 kDa protein that cross-reacts with an antibody raised against cucumber expansin. The results show that tomato leafwalls possess at least one expansin that is responsible for the acid-growth property of leaves and indicate that cell wall extension in leaves shares an underlying protein mechanism common to cell wall expansion in stems.     

Limitations to photosynthesis in coffee leaves from different canopy positions.

Limitations to photosynthesis were explored in leaves from four canopy positions of field-grown, unshaded coffee (Coffea arabica L.), a tropical tree species classified as shade-obligatory. Overall, compared to shade (lower) leaves, sun (upper) leaves had higher net carbon assimilation rate (A) (4.5 against 2.0mumolm(-2)s(-1) at most) associated with higher electron transport rate (due to a greater irradiance availability) but unrelated to stomatal and mesophyll conductances, which were similar regardless of leaf position. Neither physiological variable directly involved with photosynthetic carbon gain nor those involved with light capture were able to adjust themselves to match the capacity of the photosynthetic machinery to the light supply. We concluded that: (i) there was no major difference in photosynthetic capacity between sun and shade leaves; (ii) the intrinsic low A in coffee was greatly associated with remarkable low diffusive limitations rather than with biochemical or photochemical constraints; and (iii) morphological (e.g., variations in specific leaf area and leaf inclination) or anatomical plasticity should be of greater acclimative value than physiological plasticity as a mean of coffee leaves to respond to changing irradiance.     

Response of soybean photosynthesis and chloroplast membrane function to canopy development and mutual shading

The effect of natural shading on photosynthetic capacity and chloroplast thylakoid membrane function was examined in soybean (Glycine max. cv Young) under field conditions using a randomized complete block design. Seedlings were thinned to 15 plants per square meter at 20 days after planting. Leaves destined to function in the shaded regions of the canopy were tagged during early expansion at 40 days after planting. To investigate the response of shaded leaves to an increase in available light, plants were removed from certain plots at 29 or 37 days after tagging to reduce the population from 15 to three plants per square meter and alter the irradiance and spectral quality of light. During the transition from a sun to a shade environment, maximum photosynthesis and chloroplast electron transport of control leaves decreased by two- to threefold over a period of 40 days followed by rapid senescence and abscission. Senescence and abscission of tagged leaves were delayed by more than 4 weeks in plots where plant populations were reduced to three plants per square meter. Maximum photosynthesis and chloroplast electron transport activity were stabilized or elevated in response to increased light when plant populations were reduced from 15 to three plants per square meter. Several chloroplast thylakoid membrane components were affected by light environment. Cytochrome f and coupling factor protein decreased by 40% and 80%, respectively, as control leaves became shaded and then increased when shaded leaves acclimated to high light. The concentrations of photosystem I (PSI) and photosystem II (PSII) reaction centers were not affected by light environment or leaf age in field grown plants, resulting in a constant PSII/PSI ratio of 1.6 ± 0.3. Analysis of the chlorophyll-protein composition revealed a shift in chlorophyll from PSI to PSII as leaves became shaded and a reversal of this process when shaded leaves were provided with increased light. These results were in contrast to those of soybeans grown in a growth chamber where the PSII/PSI ratio as well as cytochrome f and coupling factor protein levels were dependent on growth irradiance. To summarize, light environment regulated both the photosynthetic characteristics and the timing of senescence in soybean leaves grown under field conditions.     

水分对番茄不同叶龄叶片光合作用的影响

以番茄品种"金棚1号"为材料,采用盆栽方式,按照蒸腾蒸发量(ET)的50%、75%、100%和125%作为补充灌溉量研究了不同水分下番茄结果期叶片气体交换特性和光响应特征参数随叶龄的变化。结果表明:番茄叶片随着叶龄的增加,净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)逐渐降低,水分利用效率(WUE)呈先上升后下降趋势;叶龄为18 d和29 d的叶片最大净光合速率(Pmax)随灌溉量的增加均先增加后降低,分别在75%ET和100%ET处理达到最大值。叶龄为38 d和47 d的叶片Pmax均以125%ET处理最大。表观量子效率(α)随叶龄的增大也先升高后下降,在叶龄为38 d时最大;番茄叶片的光饱和点(LSP)随叶龄的加大而减小。不同水分处理下不同叶龄叶片的光响应特征参数为:叶片在叶龄为18 d时,Pmax为20.64—26.73μmol.m-.2s-1,α为0.0518—0.0556;叶龄为29 d时,Pmax为11.00—24.24μmol.m-.2s-1,α为0.0522—0.0594;叶龄为38 d时,Pmax为11.77—18.18μmol.m-.2s-1,α为0.0619—0.0693;叶龄为47 d时,Pmax为9.09—18.17μmol.m-.2s-1,α为0.0538—0.0606。随叶龄加大,增加补充灌溉量有利于延缓叶片光合能力的降低。气孔限制是水分影响番茄叶片光合作用的主要因素,气孔限制与非气孔限制因素是番茄叶片Pn随叶龄变化的原因。     

遮阴对高山杜鹃叶片解剖和光合特性的影响

为了解高山杜鹃对光能的需求和适应性,该研究以盆栽3 a生高山杜鹃品种cv. Furnivall's Daughter为材料,探讨了遮阴对高山杜鹃叶片解剖结构和光合特性的影响。结果表明:光照强度对高山杜鹃品种cv. Furnivall's Daughter叶片的气孔密度没有显著影响,其气孔密度范围在299.70~327.22个·mm~(-2)之间,但光照对气孔开度和单个气孔器的面积影响显著,100%全光照和30%全光照处理植株分别具有最小和最大的叶片气孔开度。在处理的光强范围内,随着光强减弱,叶片厚度、栅栏组织厚度、海绵组织厚度以及上、下表皮厚度逐渐降低,有利于提高叶片的光能利用效率。100%全光照处理下,高山杜鹃叶片的光饱和点(LSP)、净光合速率(P_n)、饱和光合速率(P_(max))、气孔导度(Gs)、蒸腾速率(Tr)均较低,遮阴处理有效提高了Pn、P_(max)、G_s、T_r和光能利用效率(LUE),且30%全光照处理植株的叶片光补偿点(LCP)、暗呼吸速率(Rd)最低,而LSP、P_n、P_(max)、G_s、T_r和LUE最高。这表明高山杜鹃在云南昆明地区的最适光照条件是30%左右的全光照,在高山杜鹃的栽培及应用中,应采取适当的遮阴措施以满足其生长的最佳光照条件。     

Acclimation of photosynthesis to nitrogen deficiency in Phaseolus vulgaris

Nitrogen deficiency diminishes consumption of photosynthates in anabolic metabolism. We studied adjustments of the photosynthetic machinery in nitrogen-deficient bean plants and found four phenomena. First, the number of chloroplasts per cell decreased. Chloroplasts of nitrogen starved leaves contained less pigments than those of control leaves, but the in vitro activities of light reactions did not change when measured on chlorophyll basis. Second, nitrogen deficiency induced cyclic electron transfer. The amounts of Rubisco and ferredoxin-NADP⁺ reductase decreased in nitrogen starved plants. Low activities of these enzymes are expected to lead to increase in reduction of oxygen by photosystem I. However, diaminobenzidine staining did not reveal hydrogen peroxide production in nitrogen starved plants. Measurements of far-red-light-induced redox changes of the primary donor of photosystem I suggested that instead of producing oxygen radicals, nitrogen starved plants develop a high activity of cyclic electron transport that competes with oxygen for electrons. Nitrogen starvation led to decrease in photochemical quenching and increase in non-photochemical quenching, indicating that cyclic electron transport reduces the plastoquinone pool and acidifies the lumen. A third effect is redistribution of excitation energy between the photosystems in favor of photosystem I. Thus, thylakoids of nitrogen starved plants appeared to be locked in state 2, which further protects photosystem II by decreasing its absorption cross-section. As a fourth response, the proportion of non-Q_B-reducing photosystem II reaction centers increased and the redox potential of the Q_B/Q_B⁻ pair decreased by 25 mV in a fraction of photosystem II centers of nitrogen starved plants.     

Acclimation to long-term water deficit in the leaves of two sunflower hybrids: photosynthesis, electron transport and carbon metabolism

The influence of long-term water deficit on photosynthesis, electron transport and carbon metabolism of sunflower leaves has been examined. Water deficit was imposed from flower bud formation up to the stage of full flowering in the field on two sunflower hybrids with different drought tolerance. CO2 assimilation and stomatal conductance of the intact leaves, determined at atmospheric CO2 and full sunlight (1500-2000 mol quanta m^-2 s^-1), decreased with water deficit. Maximum quantum efficiency of PSII (Fy/Fm) and relative quantum yield of PSII (II) determined under similar experimental conditions, did not change significantly in severely stressed leaves. The strong inhibition of the plateau region of the light response curve, determined at high CO2 (5%) in water-deficient sunflower leaves, indicates that photosynthesis is also limited by non-stomatal factors. The decreased slope and the plateau of the CO2 response curves show that the capacity of carboxylation and RuBP regeneration decreased in severely stressed intact leaves. Rubisco specific activity decreased in severely stressed leaves, but Rubisco content increased under prolonged drought. The increase of Rubisco content was significantly higher in leaves of the drought-tolerant sunflower hybrid indicating that a higher Rubisco content could be one factor in conferring better acclimation and higher drought tolerance.     

Acclimation responses of mature Abies amabilis sun foliage to shading

This paper addresses two main questions. First, can evergreen foliage that has been structurally determined as sun foliage acclimate physiologically when it is shaded? Second, is this acclimation independent of the foliage ageing process and source-sink relations? To investigate these questions, a shading and debudding experiment was established using paired branches on opengrown Abies amabilis trees. For each tree, one branch was either shaded, debudded, or both, from before budbreak until the end of summer, while the other branch functioned as a control. Foliage samples were measured both prior to and during treatment for photosynthesis at light saturation (A _max), dark respiration, nitrogen content, chlorophyll content, chlorophyll-to-nitrogen ratio and chlorophyll a:b ratio. All age classes of foliage responded similarly during the treatment, although pre-treatment values differed between age classes. Within 1 month after the treatment began, A _max was lower in shaded foliage and remained lower throughout the treatment period. For debudded branches, A _max was lower than the controls only during active shoot elongation. At the end of the treatments in September, A _max in shade-treated sun foliage matched the rates in the true shade-formed foliage, but nitrogen remained significantly higher. By 1.5 months after treatment, chlorophyll content in shaded foliage was higher than in controls, and the chlorophyll a:b ratio was lower for the shaded foliage. On debudded branches, chlorophyll content and chlorophyll a:b ratio were similar to the values in control samples. Shading lowered the rate of nitrogen accumulation within a branch, while removing debudding decreased the amount of sequestered N that was exported from the older foliage to supply new growth. By September, chlorophyll content in shade-treated foliage was higher than that in the control sun foliage or in true shade foliage. The chlorophyll increase as a result of shading was unexpected. However, the chlorophyll-to-nitrogen ratio was identical for the shade-treated sun foliage and the true shade foliage while being significantly lower than the control sun foliage. It appears that acclimation to shading in mature foliage involves a reallocation of nitrogen within the leaf into thylakoid proteins. A redistribution of resources (nitrogen) among leaves is secondary and appears to function on a slower time scale than reallocation within the leaf. Thus, A. amabilis foliage that is structurally determined as sun foliage can acclimate to shade within a few months; this process is most likely independent of ageing and is only slightly affected by source-sink relations within a branch.     

Acclimation of photosynthesis to low leaf water potentials

Photosynthesis is reduced at low leaf water potentials (Ψ_l) but repeated water deficits can decrease this reduction, resulting in photosynthetic acclimation. The contribution of the stomata and the chloroplasts to this acclimation is unknown. We evaluated stomatal and chloroplast contributions when soil-grown sunflower (Helianthus annuus L.) plants were subjected to water deficit pretreatments for 2 weeks. The relationship between photosynthesis and Ψ_l, determined from gas-exchange and isopiestic thermocouple psychometry, was shifted 3 to 4 bars towards lower Ψ_l, in pretreated plants. Leaf diffusive resistance was similarly affected. Chloroplast activity, demonstrated in situ with measurements of quantum yield and the capacity to fix CO₂ at all partial pressures of CO₂, and in vitro by photosystem II activity of isolated organelles, was inhibited at low Ψ_l but less in pretreated plants than in control plants. The magnitude of this inhibition indicated that decreases in chloroplast activity contributed more than closure of stomata both to losses in photosynthesis and to the acclimation of photosynthesis to low Ψ_l.     

Adjustment of leaf photosynthesis to shade in a natural canopy: rate parameters

The present study was performed to investigate the adjustment of the rate parameters of the light and dark reactions of photosynthesis to the natural growth light in leaves of an overstorey species, Betula pendula Roth, a subcanopy species, Tilia cordata P. Mill., and a herb, Solidago virgaurea L., growing in a natural plant community in Järvselja, Estonia. Shoots were collected from the site and individual leaves were measured in a laboratory applying a standardized routine of kinetic gas exchange, Chl fluorescence and 820 nm transmittance measurements. These measurements enabled the calculations of the quantum yield of photosynthesis and rate constants of excitation capture by photochemical and non-photochemical quenchers, rate constant for P700⁺ reduction via the cytochrome b₆f complex with and without photosynthetic control, actual maximum and potential (uncoupled) electron transport rate, stomatal and mesophyll resistances for CO₂ transport, K_m(CO₂) and V_m of ribulose-bisphosphate carboxylase-oxygenase (Rubisco) in vivo. In parallel, N, Chl and Rubisco contents were measured from the same leaves. No adjustment toward higher quantum yield in shade compared with sun leaves was observed, although relatively more N was partitioned to the light-harvesting machinery in shade leaves ( H. Eichelmann et al., 2004 ). The electron transport rate through the Cyt b₆f complex was strongly down-regulated under saturating light compared with darkness, and this was observed under atmospheric, as well as saturating CO₂ concentration. In vivo V_m measurements of Rubisco were lower than corresponding reported measurements in vitro, and the k_cat per reaction site varied widely between leaves and growth sites. The correlation between Rubisco V_m and the photosystem I density was stronger than between V_m and the density of Rubisco active sites. The results showed that the capacity of the photosynthetic machinery decreases in shade-adjusted leaves, but it still remains in excess of the actual photosynthetic rate. The photosynthetic control systems that are targeted to adjust the photosynthetic rate to meet the plant's needs and to balance the partial reactions of photosynthesis, down-regulate partial processes of photosynthesis: excess harvested light is quenched non-photochemically; excess electron transport capacity of Cyt b₆f is down-regulated by ΔpH-dependent photosynthetic control; Rubisco is synthesized in excess, and the number of activated Rubisco molecules is controlled by photosystem I-related processes. Consequently, the nitrogen contained in the components of the photosynthetic machinery is not used at full efficiency. The strong correlation between leaf nitrogen and photosynthetic performance is not due to the nitrogen requirements of the photosynthetic apparatus, but because a certain amount of energy must be captured through photosynthesis to maintain this nitrogen within a leaf.     

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.     

Acclimation of the respiration/photosynthesis ratio to temperature: insights from a model

Based on short-term experiments, many plant growth models – including those used in global change research – assume that an increase in temperature stimulates plant respiration (R) more than photosynthesis (P), leading to an increase in the R/P ratio. Longer-term experiments, however, have demonstrated that R/P is relatively insensitive to growth temperature. We show that both types of temperature response may be reconciled within a simple substrate-based model of plant acclimation to temperature, in which respiration is effectively limited by the supply of carbohydrates fixed through photosynthesis. The short-term, positive temperature response of R/P reflects the transient dynamics of the nonstructural carbohydrate and protein pools; the insensitivity of R/P to temperature on longer time-scales reflects the steady-state behaviour of these pools. Thus the substrate approach may provide a basis for predicting plant respiration responses to temperature that is more robust than the current modelling paradigm based on the extrapolation of results from short-term experiments. The present model predicts that the acclimated R/P depends mainly on the internal allocation of carbohydrates to protein synthesis, a better understanding of which is therefore required to underpin the wider use of a constant R/P as an alternative modelling paradigm in global change research.     

Adjustment of leaf photosynthesis to shade in a natural canopy: reallocation of nitrogen

The present study was performed to investigate the adjustment of the constituents of the light and dark reactions of photosynthesis to the natural growth irradiance in the leaves of an overstorey species, Betula pendula Roth, a subcanopy species Tilia cordata P. Mill., and a herb Solidago virgaurea L. growing in a natural plant community in Järvselja, Estonia. Shoots were collected from the site and properties of individual leaves were measured in a laboratory, by applying a routine of kinetic gas exchange and optical measurements that revealed photosystem II (PSII), photosystem I (PSI), and cytochrome b₆f densities per leaf area and the distribution of excitation (or chlorophyll, Chl) between the two photosystems. In parallel, N, Chl and ribulose-bisphosphate carboxylase-oxygenase (Rubisco) content was measured from the same leaves. The amount of N in photosynthetic proteins was calculated from the measured contents of the components of the photosynthetic machinery. Non-photosynthetic N was found as the residual of the budget. Growth in shade resulted in the decrease of leaf dry mass to a half of the DW in sun leaves in each species, but the total variation, from the top to the bottom of the canopy, was larger. Through the whole cross-section of the canopy, leaf dry weight (DW) and Rubisco content per area decreased by a factor of four, N content by a factor of three, but Chl content only by a factor of 1.7. PSII density decreased by a factor of 1.9, but PSI density by a factor of 3.2. The density of PSI adjusted to shade to a greater extent than the density of PSII. In shade, the distribution of N between the components of the photosynthetic machinery was shifted toward light-harvesting proteins at the expense of Rubisco. Non-photosynthetic N decreased the most substantially, from 54% in the sun leaves of B. pendula to 11% in the shade leaves of T. cordata. It is concluded that the redistribution of N toward light-harvesting Chl proteins in shade is not sufficient to keep the excitation rate of a PSII centre invariant. Contrary to PSII, the density of PSI – the photosystem that is in immediate contact with the carbon assimilation system – shade-adjusts almost proportionally with the latter, whereas its Chl antenna correspondingly increases. Even under N deficiency, a likely condition in the natural plant community, a substantial part of N is stored in non-photosynthetic proteins under abundant irradiation, but much less under limiting irradiation. At least in trees the general sequence of down-regulation due to shade adjustment is the following: (1) non-protein cell structures and non-photosynthetic proteins; (2) carbon assimilation proteins; (3) light reaction centre proteins, first PSI; and (4) chlorophyll-binding proteins.     

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.     

Nitrogen use efficiency in instantaneous and daily photosynthesis of leaves in the canopy of a Solidago altissima stand

Photosynthetic capacity was measured on detached leaves sampled in a canopy of Solidago altissima L. Non-rectangular hyperbola fitted the light response curve of photosynthesis and significant correlations were observed between leaf nitrogen per unit area and four parameters which characterize the light-response curve. Using regressions of the parameters on leaf nitrogen, a model of leaf photosynthesis was constructed which gave the relationships between leaf nitrogen, photon flux density (PFD) and photosynthesis. Curvilinear relations were obtained between leaf nitrogen and photosynthetic rate on both an instantaneous and a daily basis. Nitrogen use efficiency (NUE, photosynthesis per unit leaf nitrogen) was calculated against leaf nitrogen under varying PFDs. The optimum nitrogen content per unit leaf area that maximizes NUE shifted to higher values with increasing PFD. Field measurements of PFD showed high positive correlations between the distribution of leaf nitrogen in the canopy and relative PFD. The predicted optimum leaf nitrogen content for each level in the canopy, to achieve maximized NUE during a clear day, was close to the actual nitrogen distribution as found through sampling.