Acclimation of photosynthetic capacity to irradiance in tree canopies in relation to leaf nitrogen concentration and leaf mass per unit area

The observation of acclimation in leaf photosynthetic capacity to differences in growth irradiance has been widely used as support for a hypothesis that enables a simplification of some soil‐vegetation‐atmosphere transfer (SVAT) photosynthesis models. The acclimation hypothesis requires that relative leaf nitrogen concentration declines with relative irradiance from the top of a canopy to the bottom, in 1 : 1 proportion. In combination with a light transmission model it enables a simple estimate of the vertical profile in leaf nitrogen concentration (which is assumed to determine maximum carboxylation capacity), and in combination with estimates of the fraction of absorbed radiation it also leads to simple ‘big‐leaf’ analytical solutions for canopy photosynthesis. We tested how forests deviate from this condition in five tree canopies, including four broadleaf stands, and one needle‐leaf stand: a mixed‐species tropical rain forest, oak (Quercus petraea (Matt.) Liebl), birch (Betula pendula Roth), beech (Fagus sylvatica L.) and Sitka spruce (Picea sitchensis (Bong.) Carr). Each canopy was studied when fully developed (mid‐to‐late summer for temperate stands). Irradiance (Q, µmol m^?2 s^?1) was measured for 20 d using quantum sensors placed throughout the vertical canopy profile. Measurements were made to obtain parameters from leaves adjacent to the radiation sensors: maximum carboxylation and electron transfer capacity (V_a, J_a, µmol m^?2 s^?1), day respiration (R_da, µmol m^?2 s^?1), leaf nitrogen concentration (N_m, mg g^?1) and leaf mass per unit area (L_a, g m^?2). Relative to upper‐canopy values, V_a declined linearly in 1 : 1 proportion with N_a. Relative V_a also declined linearly with relative Q, but with a significant intercept at zero irradiance (P < 0·01). This intercept was strongly related to L_a of the lowest leaves in each canopy (P < 0·01, r² = 0·98, n= 5). For each canopy, daily lnQ was also linearly related with lnV_a(P < 0·05), and the intercept was correlated with the value for photosynthetic capacity per unit nitrogen (PUN: V_a/N_a, µmol g^?1 s^?1) of the lowest leaves in each canopy (P < 0·05). V_a was linearly related with L_a and N_a(P < 0·01), but the slope of the V_a : N_a relationship varied widely among sites. Hence, whilst there was a unique V_a : N_a ratio in each stand, acclimation in V_a to Q varied predictably with L_a of the lowest leaves in each canopy. The specific leaf area, L_m(cm² g^?1), of the canopy‐bottom foliage was also found to predict carboxylation capacity (expressed on a mass basis; V_m, µmol g^?1 s^?1) at all sites (P < 0·01). These results invalidate the hypothesis of full acclimation to irradiance, but suggest that L_a and L_m of the most light‐limited leaves in a canopy are widely applicable indicators of the distribution of photosynthetic capacity with height in forests.     

热带雨林三种树苗叶片光合机构对光强的适应

对生长在不同光强(自然日光的8％,25％,50％)下西双版纳热带雨林3种木本植物团花(Anthocephalus chinensis)、玉蕊(Barringtonia pendala)和藤黄(Garrcinia hanburyi)幼苗光合机构的研究表明,随着生长光强的升高,植物叶片的光饱和点、补偿点、净光合速率和非光化学淬灭系数(NPQ)升高,而表现量子效率(AQY)、有效光化学量子产量(Fv／Fm)、光化学淬灭系数(qP)下降．在抗氧化系统中,超氧化物歧化酶(SOD)、抗坏血酸过氧化酶(APX)活性随着光强的升高而升高,而过氢化物酶(CAT)活性与生长光强的变化不一致．抗坏血酸(AsA)含量随着光强的升高而急剧上升。最能反映PFD的变化．可以认为,除与叶黄素循环有关的热耗散增大之外,植物叶片抗氧化系统的加强也是响应强光的一种保护措施．     

湛江5种红树林树种光合作用特性及光合固碳能力研究

应用Li-6400便携式光合作用测定系统,对湛江市特呈岛5种红树林树种的净光合速率日变化和光合作用—光响应曲线进行了测定,探讨了各树种的光合作用特性以及主要影响因子并评估其固碳能力大小。结果表明:在自然光照条件下,秋茄和红海榄叶片净光合速率(Pn)的日变化曲线呈单峰型,白骨壤、木榄和桐花树为双峰型,光合"午休"现象明显,而且峰值分别出现在10:00和14:00左右。其中,白骨壤和木榄的光合午休主要由气孔限制因素引起,桐花树主要由非气孔限制因素引起。通径分析表明,光合有效辐射(PAR)是影响白骨壤和桐花树叶片Pn的主要决策因子,而叶面大气蒸汽压亏缺(VPD)是主要限制因子;影响秋茄和红海榄叶片Pn的主要决策因子是气孔导度(Gs),主要限制因子是叶温(Tl);影响木榄叶片Pn的主要决策因子是气孔导度(Gs)。基于叶片净光合作用速率的各树种日净固碳量存在显著性差异(P0.01),秋茄的日净固碳量最大(13.83 g·m-2·d-1),其次为白骨壤和桐花树(9.48和8.24 g·m-2·d-1),木榄和红海榄的较小(6.72和6.30 g·m-2·d-1)。5种红树林树种的光补偿点(LCP)介于28.3~137.0μmol·m-2·s-1之间,显示了阳生植物的特性。光饱和点(LSP)介于169.3~1189.3μmol·m-2·s-1之间,桐花树最大,红海榄最小。5种红树林树种的表观量子效率(AQY)存在极显著差异(P0.01),白骨壤最高为0.064 mol·mol-1,木榄最低,仅为0.005 mol·mol-1。5种红树林植物叶片的光响应参数与日净固碳量的关联度大小顺序为最大净光合速率(Pmax)、LSP-LCP、AQY、LSP、LCP。     

Predicting habitat distribution and frequency from plant species co-occurrence data

Aim Species frequency data have been widely used in nature conservation to aid management decisions. To determine species frequencies, information on habitat occurrence is important: a species with a low frequency is not necessarily rare if it occupies all suitable habitats. Often, information on habitat distribution is available for small geographic areas only. We aim to predict grid‐based habitat occurrence from grid‐based plant species distribution data in a meso‐scale analysis. Location The study was carried out over two spatial extents: Germany and Bavaria. Methods Two simple models were set up to examine the number of characteristic plant species needed per grid cell to predict the occurrence of four selected habitats (species data from FlorKart, http://www.floraweb.de ). Both models were calibrated in Bavaria using available information on habitat distribution, validated for other federal states, and applied to Germany. First, a spatially explicit regression model (generalized linear model (GLM) with assumed binomial error distribution of response variable) was obtained. Second, a spatially independent optimization model was derived that estimated species numbers without using spatial information on habitat distribution. Finally, an additional uncalibrated model was derived that calculated the frequencies of 24 habitats. It was validated using NATURA2000 habitat maps. Results Using the Bavarian models it was possible to predict habitat distribution and frequency from the co‐occurrence of habitat‐specific species per grid cell. As the model validations for other German federal states were successful, the models were applied to all of Germany, and habitat distribution and frequencies could be retrieved for the national scale on the basis of habitat‐specific species co‐occurrences per grid cell. Using the third, uncalibrated model, which includes species distribution data only, it was possible to predict the frequencies of 24 habitats based on the co‐occurrence of 24% of formation‐specific species per grid cell. Predicted habitat frequencies deduced from this third model were strongly related to frequencies of NATURA2000 habitat maps. Main conclusions It was concluded that it is possible to deduce habitat distributions and frequencies from the co‐occurrence of habitat‐specific species. For areas partly covered by habitat mappings, calibrated models can be developed and extrapolated to larger areas. If information on habitat distribution is completely lacking, uncalibrated models can still be applied, providing coarse information on habitat frequencies. Predicted habitat distributions and frequencies can be used as a tool in nature conservation, for example as correction factors for species frequencies, as long as the species of interest is not included in the model set‐up.     

Contrasting patterns of photosynthetic acclimation to the light environment are dependent on the differential expression of the responses to altered irradiance and spectral quality

Chl, chlorophyll
Chl a/b, ratio of chlorophyll a to chlorophyll b
Cyt f, cytochrome f
FR, far-red light
LFR, low irradiance, far-red enriched growth light
LHCII, light harvesting complex associated with PSII
LW, low irradiance, white growth light
MW, moderate irradiance, white growth light
PAR, photosynthetically active radiation
P_max, light and CO₂ saturated photosynthetic rate
PSI, photosystem I
PSII, photosystem II

Four plant species (Chamerion angustifolium, Digitalis purpurea, Brachypodium sylvaticum and Plantago lanceolata) which have previously been shown to demonstrate contrasting photosynthetic acclimatory responses to the light environment ( 33 , Plant, Cell and Environment 20, pp. 438–448) were analysed at a biochemical level. Plants were grown under low irradiance with a shade-type spectrum (LFR: 50μmol quanta m^–2 s^–1), moderately high white light (MW: 300μmol quanta m^–2 s^–1) and low irradiance white light (LW: 50μmol quanta m^–2 s^–1). The effects of light quality upon chlorophyll content and photosynthetic capacity were found to be species-dependent. A far-red dependent reduction in chlorophyll was found in three species, and an irradiance-dependent reduction was found in B. sylvaticum, which showed the greatest alteration in the xanthophyll cycle pool size of all species tested under these conditions. Chlorophyll a/b ratios were sensitive to both light quality and quantity in C. angustifolium and D. purpurea, being highest in MW, lowest in LFR, and intermediate in LW, whilst the other species showed no response. Ratios of photosystem II to photosystem I (PSII and PSI) demonstrated a strong irradiance-associated increase in all species except B. sylvaticum, whereas an increase in PSII/PSI in LFR compared to LW conditions was present in all species. A change in chlorophyll a/b was not always associated with a change in PSII/PSI, suggesting that the level of LHCII associated with each PSII varied in some species. Cytochrome f content showed an irradiance-dependent effect only, indicating a relationship with the capacity of electron transport. It is concluded that differing strategies of acclimation to the light environment demonstrated by these species results from differing strengths of expression of a series of independently regulated changes in the levels of photosynthetic components.     

Small-scale heterogeneity in soil quality influences photosynthetic efficiency and habitat selection in a clonal plant

BACKGROUND AND AIMS: In clonal plants, internode connections allow translocation of photosynthates, water, nutrients and other substances among ramets. Clonal plants form large systems that are likely to experience small-scale spatial heterogeneity. Physiological and morphological responses of Fragaria vesca to small-scale heterogeneity in soil quality were investigated, together with how such heterogeneity influences the placement of ramets. As a result of their own activities plants may modify the suitability of their habitats over time. However, most experiments on habitat selection by clonal plants have not generally considered time as an important variable. In the present study, how the foraging behaviour of clonal plants may change over time was also investigated. METHODS: In a complex of environments with different heterogeneity, plant performance was determined in terms of biomass, ramet production and photosynthetic activity. To identify habitat selection, the number of ramets produced and patch where they rooted were monitored. KEY RESULTS: Parent ramets in heterogeneous environments showed significantly higher maximum and effective quantum yields of photosystem II than parents in homogeneous environments. Parents in heterogeneous environments also showed significantly higher investment in photosynthetic biomass and stolon/total biomass, produced longer stolons, and had higher mean leaf size than parents in homogeneous environments. Total biomass and number of offspring ramets were similar in both environments. However, plants in homogeneous environments showed random allocation of offspring ramets to surrounding patches, whereas plants in heterogeneous environments showed preferential allocation of offspring to higher-quality patches. CONCLUSIONS: The results suggest that F. vesca employs physiological and morphological strategies to enable efficient resource foraging in heterogeneous environments and demonstrate the benefits of physiological integration in terms of photosynthetic efficiency. The findings indicate that short-term responses cannot be directly extrapolated to the longer term principally because preferential colonization of high-quality patches means that these patches eventually show reduced quality. This highlights the importance of considering the time factor in experiments examining responses of clonal plants to heterogeneity.     

西鄂尔多斯4种荒漠植物光合作用特征与差异性

植物对光能的高效吸收、传递和转换机理是光合作用的核心。为了厘清西鄂尔多斯地区4种荒漠植物光合生理生态适应性和生境适宜性,运用LI-6800光合作用测定系统对四合木(Tetraena mongolica)、霸王(Sarcozygium xanthoxylon)、沙冬青(Ammopiptanthus mongolicus)和白刺(Nitraria tangutorum)进行光合作用日变化进行了测定。研究结果表明:(1) 4种荒漠植物的光合作用日变化的净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)和胞间CO_2浓度(Ci)均表现出明显的分异;(2)净光合速率均呈"双峰"曲线的变化趋势,4种荒漠植物的净光合速率(Pn)依次排序为四合木(4.37μmol m~(-2) s~(-1))霸王(3.58μmol m~(-2) s~(-1))沙冬青(2.63μmol m~(-2) s~(-1))和白刺(2.26μmol m~(-2) s~(-1))。说明四合木和近缘种霸王比其他二种荒漠植物具有较强的光合生理生态适应性与生境适宜性;(3)从影响光合作用的有关生理生态因子来看,净光合速率与气孔导度与蒸腾速率、水分利用效率呈现明显的正相关性,与微气象因子大气相对湿度(RH)的相关性不明显;(4)水分因子是限制4种荒漠植物生长的重要因素之一。该研究以期为我国西北荒漠区珍稀濒危植物的保护提供案例借鉴与理论依据。     

Acclimation of photosynthesis and respiration is asynchronous in response to changes in temperature regardless of plant functional group

Gas exchange, fluorescence, western blot and chemical composition analyses were combined to assess if three functional groups (forbs, grasses and evergreen trees/shrubs) differed in acclimation of leaf respiration (R) and photosynthesis (A) to a range of growth temperatures (7, 14, 21 and 28 degrees C). When measured at a common temperature, acclimation was greater for R than for A and differed between leaves experiencing a 10-d change in growth temperature (PE) and leaves newly developed at each temperature (ND). As a result, the R : A ratio was temperature dependent, increasing in cold-acclimated plants. The balance was largely restored in ND leaves. Acclimation responses were similar among functional groups. Across the functional groups, cold acclimation was associated with increases in nonstructural carbohydrates and nitrogen. Cold acclimation of R was associated with an increase in abundance of alternative and/or cytochrome oxidases in a species-dependent manner. Cold acclimation of A was consistent with an initial decrease and subsequent recovery of thylakoid membrane proteins and increased abundance of proteins involved in the Calvin cycle. Overall, the results point to striking similarities in the extent and the biochemical underpinning of acclimation of R and A among contrasting functional groups differing in overall rates of metabolism, chemical composition and leaf structure.     

Acclimation to irradiance in seedlings of three tropical rain forest Garcinia species after simulated gap formation

We investigated the acclimation of seedlings of three tropical rain forest sub-canopy Garcinia species (G. xanthochymus, G. cowa, and G. bracteata) after transfer from 4.5 (LI) to 40 % (HI) sunlight and 12.5 (MI) sunlight to HI (LH₁ and LH₂ denoting transfer from LI to HI and MI to HI transfer, respectively). The changes of chlorophyll (Chl) fluorescence, net photosynthetic rate (P _N), dark respiration rate (R _D), Chl content per unit area (Chl_area), leaf mass per unit area (LMA), and seedling mortality were monitored over two months after transfer. These parameters together with leaf anatomy of transferred and control seedlings (kept in LI, MI, and HI) were also examined after two months. No seedlings died during the two months. F_v/F_m, P _N, and Chl_area of the transferred seedlings decreased in the first 3 to 12 d. LH₁ leaves showed larger reduction in F_v/F_m (>23 % vs. <16 %) and slower recovery of F_v/F_m than LH₂ leaves. P _N started to recover after about one week of I transfer and approached higher values in all G. cowa seedlings and G. xanthochymus LH₁ seedlings than those before the transfer. However, P _N of G. bracteata seedlings approached the values before transfer. The final P _N values in leaves of transferred G. xanthochymus and G. cowa seedlings approached that of leaves kept in HI, while the final P _N values of transferred leaves of G. bracteata were significantly lower than that of leaves grown under HI (p<0.05). R _D of G. xanthochymus LH₁ seedlings and all G. cowa seedlings increased and approached the value of the seedlings in HI. The final Chl_area of both G. xanthochymus and G. cowa approached the values before transfer, but that of G. bracteata did not recover to the level before transfer. The final Chl_area of all transferred seedlings was not significantly different from that of seedlings in HI except that G. cowa LH₁ seedlings had higher Chl_area than that in HI. LMA decreased within 2 d and then increased continuously until about 30 d and approached the value under HI. Spongy/palisade mesophyll ratio decreased after transfer because of the increase in palisade thickness. Leaf thickness did not change, so LMA increase of transferred seedlings was mainly due to the increase of leaf density. Thus the mature leaves under LI and MI of G. xanthochymus and G. cowa are able to acclimate to HI by leaf physiological and anatomical adjustment, while G. bracteata had limited ability to acclimate to HI.     

Acclimation of photosynthesis to high irradiance in rice: gene expression and interactions with leaf development

Rice (Oryza sativa L.) has been used to study the long-term responses of photosynthesis to high irradiance focusing on the composition of the photosynthetic apparatus and leaf morphology. Typical sun/shade differences in chloroplast composition are seen in the fifth leaf following growth in high irradiance compared with low irradiance (1000 and 200 micromol m(-2) s(-1), respectively): higher light-saturated rates of photosynthesis (P(max)), higher amounts of Rubisco protein, and a lower chlorophyll a:b ratio. In addition, leaves were thicker under high light compared with low light. However, responses appear more complex when leaf developmental stage is considered. Using a system of transferring plants from low to high light in the laboratory responses that occur before and after full leaf extension have been studied. Acclimation of photosynthesis is limited by leaf age: the transfer to high light, post-leaf extension, is characterized by alterations in chlorophyll a:b but not in Rubisco protein, which may be limited by leaf morphology. Microarray analysis of gene expression was carried out on plants that were transferred to high light post-leaf extension. A down-regulation of light-harvesting genes was seen. No change in the expression level of Rubisco genes was observed. Up-regulation of genes involved in photoprotection was observed. It was also shown that high-light leaf morphology is established prior to formation of the zone of cellular elongation and division. The endogenous and environmental factors which establish the characteristics of high light acclimation may be important for attaining high rates of assimilation in leaves and crop canopies, and the fifth leaf in rice provides a convenient model system for the determination of the mechanisms involved.     

Habitat quality favoured over familiarity: a rejection of natal habitat preference induction in the mosquito Aedes albopictus

1. Natal habitat preference induction (NHPI) is a behavioural phenomenon in which offspring show a change in preference in adult oviposition choice as a function of experience as an immature. 2. Although well known in certain systems, such as herbivorous insects, this behaviour has not been well studied in aquatic insects. 3. The container–breeding mosquito, Aedes albopictus (Skuse) was used to test if NHPI occurs in aquatic insects under natural conditions of two leaf species as a nutritive base (Juniperus virginiana L. and Quercus virginiana Mill) and two larval densities. 4. Significant effects of leaf species and density on adult mosquito attributes were found, with J. virginiana and low larval density associated with more, faster developing, larger and more fecund mosquitoes. However, no evidence for NHPI was found. Instead a canalised behavior was found that included spreading eggs between high– and low–quality oviposition choices in the same proportions regardless of larval experience.     

Habitat distribution and habitat preference in Barbary macaques (Macaca sylvanus)

Barbary macaques (Macaca sylvanus,L.) occur at varying densities in a number of different habitat types in Morocco and Algeria. Taub (1977) has argued that the abundance of the species in cedar forests of the Moyen Atlas, Morocco, is an indication of a habitat preference. A reexamination of available data on the distribution and abundance of Barbary macaques suggests that monkey numbers reflect the distribution of habitat size rather than habitat types. Differences between populations relative to habitat types can be seen only between forest and scrub localities. Human factors are considered more important in determining the present status of this eclectic feeder. The modern distribution of the Barbary macaque is inadequate evidence for a habitat preference.     

Oscillations in photosynthetic carbon assimilation and chlorophyll fluorescence are different in Amaranthus caudatus,a C4 plant,and Spinacia oleracea,a C3 plant

The characteristics of oscillations in photosynthetic carbon fixation and chlorophyll fluorescence in leaves of the C₄ plant Amaranthus caudatus L. were compared to those shown by the C₃ plant Spinacia oleracea L. As in spinach, oscillations could be observed in Amaranthus when leaves were illuminated after periods of darkening, particularly at temperatures below 20°C, less so or not at all at higher temperatures. However, in contrast to spinach, pronounced oscillations occurred in Amaranthus after a sudden dark/light transition only at low, not at high photon flux densities. Whereas in spinach maxima in carbon uptake were observed slightly after minima in chlorophyll fluorescence had occurred, in Amaranthus maxima in carbon uptake were close to maxima in chlorophyll fluorescence. Since the quantum efficiency of electron transport through photosystem II of the chloroplast electron-transport chain was higher during the minima of chlorophyll fluorescence than during the maxima, the observations suggest that in Amaranthus photosynthetic water oxidation did not occur as synchronously with carbon uptake as in spinach. It is proposed that, in contrast to spinach, photosynthetic oscillations in Amaranthus are related to the diffusional transport of photosynthetic intermediates between mesophyll and bundle-sheath cells.Abbreviations F_o, F_m, F_s initial, maximal and steady-state chlorophyll a fluorescence - PFD photon flux density - Q_A primary quinone acceptor of PSII We are grateful to Professors D.A. Walker, FRS, Robert Hill Institute, University of Sheffield, Sheffield, UK., and Agu Laisk, Chair of Plant Physiology, University of Tartu, Tartu, Estonia, for helpful discussions and to Ms. S. Neimanis for help with the experiments. Our work was performed within the research of the Sonderforschungsbereich 251 of the University of Würzburg. It was supported by the Stiftung Volkswagenwerk. A.S.R. acknowledges also support by the Alexander-von-Humboldt-Stiftung and U.G. by the Graduate College of the University of Würzburg.     

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.     

冬虫夏草与产地植物及土壤的重金属含量测定

本文测定了野生采集和人工培育冬虫夏草、人工培育冬虫夏草子实体、冬虫夏草发酵菌液、冬虫夏草寄主蝙蝠蛾昆虫食物(蕨麻Potentilla anserine和珠芽蓼Polygonum viviparum),以及野生冬虫夏草产地土壤中5种重金属元素(铜Cu、铅Pb、砷As、镉Cd、汞Hg)的含量。结果表明,人工培育冬虫夏草、人工培育冬虫夏草子实体、冬虫夏草发酵菌液、蕨麻块根、珠芽蓼种子的5种重金属元素含量符合《中医药-中药材重金属限量》ISO国际标准;而野生冬虫夏草,除砷含量(6.170 mg/kg)超标外,铜、铅、镉、汞元素含量均符合要求。按国家标准《土壤环境质量农用地土壤污染风险管控标准(试行)》(GB 1516-2018),四川产地土壤(KDX)与青海产地土壤(GD、HN)的5种重金属含量均低于农用土壤污染风险筛选值;而四川产地土壤(KDT),除镉含量(0.923 mg/kg)高于风险筛选值而低于风险管制值外,其余4种元素含量均低于风险筛选值。结果为人工培育冬虫夏草的质量控制提供了参考。     

Effects of shade treatments on the photosynthetic capacity,chlorophyll fluorescence,and chlorophyll content of Tetrastigma hemsleyanum Diels et Gilg

Tetrastigma hemsleyanum Diels et Gilg was grown under full sunlight and moderate and high levels of shade for one month to evaluate its photosynthetic and chlorophyll fluorescence response to different light conditions. The results showed that T. hemsleyanum attained greatest leaf size and P_n when cultivated with 67% shade. Leaves of seedlings grown with 90% shade were the smallest. Leaf color of plants grown under full sunlight and 50% shade was yellowish-green. The P_n value increased rapidly as PPFD increased to 200 μmol m^?2 s^?1 and then increased slowly to a maximum, followed by a slow decrease as PPFD was increased to 1000 μmol m^?2 s^?1. P_n was highest for the 67% shade treatment and the LSP for this shade treatment was 600 μmol m^?2 s^?1. Full sunlight and 50% shade treatments resulted in significant reduction of ETR and qP and increased NPQ. Chl a, Chl b and total chlorophyll content increased and Chl a/b values decreased with increased shading. Results showed that light intensity greater than that of 50% shade depressed photosynthetic activity and T. hemsleyanum growth. Irradiance less than that of 75% shade limited carbon assimilation and led to decreased plant growth. Approximately 67% shade is suggested to be the optimum light irradiance condition for T. hemsleyanum cultivation.     

Protection of the photosynthetic apparatus against dehydration stress in the resurrection plant Craterostigma pumilum

The group of homoiochlorophyllous resurrection plants evolved the unique capability to survive severe drought stress without dismantling the photosynthetic machinery. This implies that they developed efficient strategies to protect the leaves from reactive oxygen species (ROS) generated by photosynthetic side reactions. These strategies, however, are poorly understood. Here, we performed a detailed study of the photosynthetic machinery in the homoiochlorophyllous resurrection plant Craterostigma pumilum during dehydration and upon recovery from desiccation. During dehydration and rehydration, C. pumilum deactivates and activates partial components of the photosynthetic machinery in a specific order, allowing for coordinated shutdown and subsequent reinstatement of photosynthesis. Early responses to dehydration are the closure of stomata and activation of electron transfer to oxygen accompanied by inactivation of the cytochrome b₆f complex leading to attenuation of the photosynthetic linear electron flux (LEF). The decline in LEF is paralleled by a gradual increase in cyclic electron transport to maintain ATP production. At low water contents, inactivation and supramolecular reorganization of photosystem II becomes apparent, accompanied by functional detachment of light‐harvesting complexes and interrupted access to plastoquinone. This well‐ordered sequence of alterations in the photosynthetic thylakoid membranes helps prepare the plant for the desiccated state and minimize ROS production.     

Slow response of plant species richness to habitat loss and fragmentation

We examined the response of vascular plant species richness to long-term habitat loss and fragmentation of Estonian calcareous grasslands (alvars). The current number of habitat specialist species in 35 alvars was not explained by their current areas and connectivities but it was explained by their areas and connectivities 70 years ago ( R ² = 0.27). We estimated the magnitude of extinction debt in local communities by assuming an equilibrium species richness in 14 alvars that had lost only a small amount of area and by applying this model to the remaining alvars, in which the average area has declined from 3.64 km² in the 1930s to 0.21 km² at present. The extinction debt estimated for individual alvars was around 40% of their current species number. Our conclusions are applicable to temperate grasslands in general, which have lost much area because of agricultural intensification and cessation of traditional management.     

Acclimation to irradiance of leaf photosynthesis and associated nitrogen reallocation in photosynthetic apparatus in the year following thinning of a young stand of Chamaecyparis obtusa

In order to quantify the effects of thinning on photosynthetic parameters and associated change in leaf nitrogen (N) contents, half of the trees in a 10-year-old Chamaecyparis obtusa (Sieb. et Zucc.) Endl. stand (36° 3′N, 140°7′E) were removed, giving a final density of 1 500 trees ha⁻¹, in May 2004. Photosynthetic photon flux density (PPFD) and leaf N and carbon (C) contents in the lower (L), middle (M), and upper (U) crowns were monitored one, three, and five months after thinning in both the thinned stand and a non-thinned control stand. In addition, leaves’ photosynthetic responses to CO₂ concentration were simultaneously measured in situ to estimate the maximum rates of carboxylation (V_cmax) and electron transport (J_max). Thinning increased PPFD in the L and M crowns but not in the U crown. V_cmax in both the L and M crowns of the thinned stand increased significantly in comparison with the same crown position of the control stand in the three and five months following thinning. In addition, the thinned stand exhibited an increase in N partitioned to ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) in the L and M crowns relative to the control stand three and five months after thinning, indicating that N had been redistributed within the photosynthetic machinery. Thinning did not affect N per unit area at any of the crown positions, but significantly increased the content of N as a fraction of the total leaf dry mass in the L and M crowns three and five months after thinning. This was a consequence of a decrease in leaf dry mass due to rapid shoot growth. Thus thinning did not cause a redistribution of N between leaves. Thinning improved irradiance in the L and M crowns of C. obtusa, leading to photosynthetic acclimation. Photosynthetic acclimation in the first year mainly occurred via redistribution of N within but not between leaves.