Ontogenetic changes in stomatal and biochemical limitations to photosynthesis of two co-occurring Mediterranean oaks differing in leaf life span

A quantitative analysis was applied to the stomatal and biochemical limitations to light-saturated net photosynthesis under optimal field conditions in mature trees and seedlings of the co-occurring evergreen oak, Quercus ilex L., and the deciduous oak, Q. faginea Lam. Stomatal limitation to photosynthesis, maximal Rubisco activity and electron transport rate were determined from assimilation versus intercellular leaf carbon dioxide concentration response curves of leaves that were subsequently analyzed for nitrogen (N) concentration, mass per unit area, thickness and percent internal air space. In both species, seedlings had a lower leaf mass per unit area, thickness and leaf N concentration than mature trees. The root system of seedlings during their third year after planting was dominated by a taproot. A lower leaf N concentration of seedlings was associated with lower maximal Rubisco activity and electron transport rate and with assimilation rates similar to or lower than those of mature trees, despite the higher stomatal conductances and potential photosynthetic nitrogen-use efficiencies of seedlings. Consequently, stomatal limitation to photosynthesis increased with tree age in both species. In both seedlings and mature trees, a lower assimilation rate in Q. ilex than in Q. faginea was associated with lower stomatal conductance, N allocation to photosynthetic functions, maximal Rubisco activity and electron transport rate, and potential photosynthetic nitrogen-use efficiency but greater leaf thickness and leaf mass per unit area. Tree-age-related changes differed quantitatively between species, and the characteristics of the two species were more similar in seedlings than in mature trees. Despite higher stomatal conductances, seedlings are more N limited than adult trees, which contributes to lower biochemical efficiency.     

土壤干旱胁迫及复水对紫叶矮樱光合特性的影响

以盆栽3年生紫叶矮樱为试材,采用称重控水法,研究土壤干旱胁迫及复水对紫叶矮樱光合特性的影响。结果表明：干旱条件下,随着胁迫程度的加重,紫叶矮樱的叶水势、净光合速率、蒸腾速率、气孔导度和胞间CO2浓度下降,叶绿素、花青苷含量和水分利用效率先升高再降低;复水后,叶水势、净光合速率、蒸腾速率、叶绿素含量、花青苷含量和水分利用效率升高,表明紫叶矮樱具有胁迫后补偿生长特征。     

光叶子花不同叶位叶片叶绿素含量和光合作用研究

本文以光叶子花炼苗60 d组培苗为材料,对光叶子花不同叶位叶绿素含量、呼吸作用(Re)、气孔导度(Gs)、胞间CO2浓度(Ci)、净光合速(Pn)、蒸腾速率(Tr)、叶面饱和水汽压亏缺(Vpdl)等进行了测定。试验结果表明,光叶子花叶片单位重量和单位面积叶绿素含量、净光合速度和水分利用效率(WUE)均随叶位上升呈先增大后减小趋势变化,其最大值都出现在第6叶位叶片;叶片呼吸速率、气孔导度和蒸腾速率均随叶位上升而减小;叶片胞间CO2浓度随叶位上升呈先减小后增大变化;叶面饱和水汽压亏缺随叶位上升而增大。由于光叶子花中部叶片叶绿素含量、Pn和WUE相对较高,因此在今后的栽培管理中应对中部叶片加强保护。     

Influence of environmental and plant factors on canopy photosynthesis and transpiration of apple trees

We estimated carbon and water flows, canopy conductance and the assimilation/transpiration ratio of fruiting and non-fruiting apple trees grown in the field, from daily gas exchange measurements taken during the summer with a whole-canopy enclosure device. The relationships between photosynthetic and transpirational responses and environmental conditions were also investigated, as well as the role of canopy conductance in controlling carbon dioxide and water vapor exchange. Light-saturated net photosynthetic rates, which were higher for the fruiting canopy than for the non-fruiting canopy, showed a general decrease in the afternoon, particularly for the non-fruiting canopy, compared with rates in the morning. When light was not limiting, the afternoon decrease in net photosynthesis appeared to be regulated more by non-stomatal factors than by changes in canopy conductance. Canopy conductance, which was higher for the fruiting canopy than for the non-fruiting canopy, may actively regulate photosynthetic activity and may also be modulated by feedback control in response to assimilation capacity. We conclude that adjustments in canopy conductance, which were partially dependent on the vegetative-reproductive status of the tree, control the equilibrium between photosynthesis and transpiration. We also demonstrated that whole-canopy chambers can be used to estimate photosynthetic and transpirational responses thereby overcoming the difficulty of scaling these physiological responses from the leaf to the whole-canopy level.     

黑河流域荒漠地区水分胁迫对梭梭苗木影响的试验研究(英文)

梭梭(HaloxylonAmmodendronBge,一种C4灌木)苗种植在15升的容器中,给予不同的水分胁迫处理,研究了其水分关系和气体交换特征。结果表明:当土壤水分含量大于11%时,梭梭苗有高的蒸腾量;土壤水分含量低于6%时,苗木就不能从土壤中吸取水分;很好供水的苗木的蒸腾量与潜在蒸发量成线型相关。气体交换测定发现,随着土壤水分含量的下降,造成了不同程度的气孔导度、叶蒸腾强度和光合作用的下降。对同一苗木而言,由于这个地区有高的水气压亏缺(VPD),很好和中度供水的苗木在气孔反应方面有较宽的范围,气孔在决定光合作用方面起着较小的作用,二者没有明显的线型相关关系。虽然水分胁迫使蒸腾速率比光合速率下降的更快,提高了水分利用效率,而较高的蒸发需求增加了蒸腾量,限制了光合作用,但是总的趋势是光合作用和蒸腾强度成线型相关。图6表2参15。     

Recovery of diurnal depression of leaf hydraulic conductance in a subtropical woody bamboo species: embolism refilling by nocturnal root pressure

Despite considerable investigations of diurnal water use characteristics in different plant functional groups, the research on daily water use strategies of woody bamboo grasses remains lacking. We studied the daily water use and gas exchange of Sinarundinaria nitida (Mitford) Nakai, an abundant subtropical bamboo species in Southwest China. We found that the stem relative water content (RWC) and stem hydraulic conductivity (K(s)) of this bamboo species did not decrease significantly during the day, whereas the leaf RWC and leaf hydraulic conductance (K(leaf)) showed a distinct decrease at midday, compared with the predawn values. Diurnal loss of K(leaf) was coupled with a midday decline in stomatal conductance (g(s)) and CO(2) assimilation. The positive root pressures in the different habitats were of sufficient magnitude to refill the embolisms in leaves. We concluded that (i) the studied bamboo species does not use stem water storage for daily transpiration; (ii) diurnal down-regulation in K(leaf) and gs has the function to slow down potential water loss in stems and protect the stem hydraulic pathway from cavitation; (iii) since K(leaf) did not recover during late afternoon, refilling of embolism in bamboo leaves probably fully depends on nocturnal root pressure. The embolism refilling mechanism by root pressure could be helpful for the growth and persistence of this woody monocot species.     

油茶不同叶龄叶片形态与光合参数的测定

为了探讨油茶叶片形态特征、光合特性与叶龄的相关性,从而为油茶的生产经营提供理论参考,利用CI-203叶面积分析仪和LI-6400便携式光合仪测定了油茶新梢不同叶龄叶片的形态特征和光合参数。结果表明:油茶叶面积和叶片鲜质量,从第1到第3位叶逐渐变大,又从第3到第7位叶逐渐减小;叶片的叶绿素含量和净光合速率均随着叶龄的增大而增大;叶龄与蒸腾速率和气孔导度均呈极显著正相关,而与胞间CO2浓度呈极显著负相关;叶片比叶面积与胞间CO2浓度间呈极显著正相关,而与净光合速率、蒸腾速率和气孔导度间均呈极显著负相关;叶片干物质含量比值与其净光合速率、气孔导度和蒸腾速率间均呈极显著负相关;叶片生长20 d,叶面积等形态特征基本发育完全,经过60 d的生长后,基本成为主要功能叶。     

Patchy stomatal behavior during midday depression of leaf CO₂ exchange in tropical trees

We investigated effects of heterogeneous stomatal behavior on diurnal patterns of leaf gas exchange in 10 tree species. Observations were made in middle and upper canopy layers of potted tropical rainforest trees in a nursery at the Forest Research Institute Malaysia. Measurements were taken from 29 January to 3 February 2010. We measured in situ diurnal changes in net photosynthetic rate and stomatal conductance in three leaves of each species under natural light. In both top-canopy and sub-canopy species, midday depression of net assimilation rate occurred in late morning. Numerical analysis showed that patchy bimodal stomatal behavior occurred only during midday depression, suggesting that the distribution pattern of stomatal apertures (either uniform or non-uniform stomatal behavior) varies flexibly within single days. Direct observation of stomatal aperture using Suzuki's Universal Micro-Printing (SUMP) method demonstrated midday patchy stomatal closure that fits a bimodal pattern in Shorea leprosula Miq., Shorea macrantha Brandis. and Dipterocarpus tempehes V.Sl. Inhibition of net assimilation rate and stomatal conductance appears to be a response to changes in vapor pressure deficit (VPD). Variable stomatal closure with increasing VPD is a mechanism used by a range of species to prevent excess water loss from leaves through evapotranspiration (viz., inhibition of midday leaf gas exchange). Bimodal stomatal closure may occur among adjacent stomata within a single patch, rather than among patches on a single leaf. Our results suggest the?occurrence of patches at several scales within single leaves. Further analysis should consider variable spatial scales in heterogeneous stomatal behavior between and within patches and within single leaves.     

Seasonal fluctuations and temperature dependence in photosynthetic parameters and stomatal conductance at the leaf scale of Populus euphratica Oliv

A combined model to simulate CO? and H?O gas exchange at the leaf scale was parameterized using data obtained from in situ leaf-scale observations of diurnal and seasonal changes in CO? and H?O gas exchange. The Farquhar et al.-type model of photosynthesis was parameterized by using the Bayesian approach and the Ball et al.-type stomatal conductance model was optimized using the linear least-squares procedure. The results show that the seasonal physiological changes in photosynthetic parameters (e.g., V(cmax25), J(max25), R(d25) and g(m25)) in the biochemical model of photosynthesis and m in the stomatal conductance model should be counted in estimating long-term CO? and H?O gas exchange. Overall, the coupled model successfully reproduced the observed response in net assimilation and transpiration rates.     

Induction of photosynthesis and importance of limitations during the induction phase in sun and shade leaves of five ecologically contrasting tree species from the temperate zone

We examined the principal differences in photosynthetic characteristics between sun and shade foliage and determined the relative importance of biochemical and stomatal limitations during photosynthetic induction. Temperate-zone broadleaf and conifer tree species, ranging widely in shade tolerance, were investigated from one locality in the Czech Republic. The study species included strongly shade-tolerant Abies alba Mill. and Tilia cordata Mill., less shade-tolerant Fagus sylvatica L. and Acer pseudoplatanus L. and sun-demanding Picea abies (L.) Karst. In the fully activated photosynthetic state, sun foliage of all species had significantly higher maximum CO(2) assimilation rates, maximum stomatal conductance and maximum rates of carboxylation than shade foliage. Compared with shade leaves, sun leaves had significantly higher nocturnal stomatal conductances. In all species, shade foliage tended to have higher induction states 60 s after leaf illumination than sun foliage. Sun and shade foliage did not differ in the rate of disappearance of the transient biochemical limitation during the induction phase. Longer time periods were required to reach 90% photosynthetic induction and 90% stomatal induction in sun foliage than in shade foliage of the less shade-tolerant F. sylvatica and A. pseudoplatanus and in sun-demanding P. abies; however, in sun foliage of the strongly shade-tolerant species T. cordata and A. alba, the time needed for photosynthetic induction was similar to, or less than, that for shade foliage. Shade but not sun needles of P. abies and A. alba had significantly slower induction kinetics than the broadleaf tree species. Among species, the sun-demanding P. abies exhibited the shortest stomatal induction times in both sun and shade leaves. Independently of shade tolerance ranking, the transient stomatal and total limitations that characterize photosynthetic induction were relieved significantly earlier in shade foliage than in sun foliage. Sun foliage generally exhibited a hyperbolic photosynthetic induction response, whereas a sigmoidal induction response was more frequent in shade foliage. The different relative proportions of transient biochemical and stomatal limitations during photosynthetic induction in sun and shade foliage indicate an essential role of stomata in photosynthetic limitation during induction, mainly in shade foliage, with a consequent influence on the shape of the photosynthetic induction curve.     

Vertical profiles reveal impact of ozone and temperature on carbon assimilation of Betula pendula and Populus tremula

Rising temperature and tropospheric ozone (O(3)) concentrations are likely to affect carbon assimilation processes and thus the carbon sink strength of trees. In this study, we investigated the joint action of elevated ozone and temperature on silver birch (Betula pendula) and European aspen (Populus tremula) saplings in field conditions by combining free-air ozone exposure (1.2?×?ambient) and infrared heaters (ambient +1.2 °C). At leaf level measurements, elevated ozone decreased leaf net photosynthesis (P(n)), while the response to elevated temperature was dependent on leaf position within the foliage. This indicates that leaf position has to be taken into account when leaf level data are collected and applied. The ozone effect on P(n) was partly compensated for at elevated temperature, showing an interactive effect of the treatments. In addition, the ratio of photosynthesis to stomatal conductance (P(n)/g(s) ratio) was decreased by ozone, which suggests decreasing water use efficiency. At the plant level, the increasing leaf area at elevated temperature resulted in a considerable increase in photosynthesis and growth in both species.     

Seasonal CO(2) assimilation and stomatal limitations in a Pinus taeda canopy

Net CO(2) assimilation (A(net)) of canopy leaves is the principal process governing carbon storage from the atmosphere in forests, but it has rarely been measured over multiple seasons and multiple years. I measured midday A(net) in the upper canopy of maturing loblolly pine (Pinus taeda L.) trees in the piedmont region of the southeastern USA on 146 sunny days over 36 months. Concurrent data for leaf conductance and photosynthetic CO(2) response curves (A(net)-C(i) curves) were used to estimate the relative importance of stomatal limitations to CO(2) assimilation in the field. In fully expanded current-year and 1-year-old needles, midday light-saturated A(net) was constant over much of the growing season (5-6 &mgr;mol CO(2) m(-2) s(-1)), except during drought periods. During the winter season (November-March), midday A(net) of overwintering needles varied in proportion to leaf temperature. Net CO(2) assimilation at light saturation occurred when daytime air temperatures exceeded 5-6 degrees C, as happened on more than 90% of the sunny winter days. In both age classes of foliage, winter carbon assimilation accounted for approximately 15% of the daily carbon assimilation on sunny days throughout the year, and was relatively insensitive to year-to-year differences in temperature during this season. However, strong stomatal limitations to A(net) occurred as a result of water stress associated with freezing cycles in winter. During the growing season, drought-induced water stress produced the largest year-to-year differences in seasonal CO(2) assimilation on sunny days. Seasonal A(net) was more drought sensitive in current-year needles than in 1-year-old needles. Relative stomatal limitations to daily integrated A(net) were approximately 40% over the growing season, and summer drought rather than high temperatures had the largest impact on summer A(net) and integrated annual CO(2) uptake in the upper crown. Despite significant stomatal limitations, a long duration of near-peak A(net) in the upper crown, particularly in 1-year-old needles, conferred high seasonal and annual carbon gain.     

Changes in leaf water use after removal of leaf lower surface hairs on <Emphasis Type="Italic">Mallotus macrostachyus</Emphasis> (Euphorbiaceae) in a tropical secondary forest in Malaysia

Leaf hairs may assist in maintaining high leaf water use efficiency in tropical secondary forest tree species. We compared leaf temperature, transpiration, photosynthesis and water use efficiency between hairy and depilated leaves in Mallotus macrostachyus (Euphorbiaceae), to determine the role of leaf hair in leaf water use efficiency (WUE) in tropical degraded secondary forest in Malaysia. Measurements were made on five mature individuals growing in sun-exposed conditions and five in shaded conditions. The hair dry weight per unit leaf area was significantly greater in sun leaves than in shade leaves. The transpiration rate (Tr_max) of depilated leaves in sun-exposed conditions was slightly higher than in hairy leaves in both morning and afternoon measurements. In contrast, Tr_max in the shade leaves was almost identical in hairy and depilated leaves. Leaf stomatal conductance (g _s) in the morning showed almost the same value among leaf types and light conditions. In the afternoon, g _s slightly decreased from the morning values in both sun and shade conditions. In the morning, the leaf water use efficiency (A _max/Tr_max) in both conditions did not differ significantly between hairy and depilated leaves. However, in the afternoon, WUE in the depilated leaves was significantly lower than in hairy leaves in sun-exposed conditions. These observations suggest that leaf hairs in M. macrostachyus contribute to the high leaf water use efficiency in drought conditions, such as high vapor pressure deficit experienced at midday in degraded tropical secondary forests.     

Diurnal Course of Gas Exchange and Water Use Efficiency of Platycladus orientalis in the Semiarid Region of the Loess Plateau

The daily gas exchange, stomatal conductance, and water use efficiency of Platycladus orientalis in the test field of Fangshan county of Shanxi Province in the semiarid region of the Loess Plateau were measured with portable Li-6200 gas analysis system in natural conditions. The results showed that the diurnal course of net photosynthetic rate displayed a two-peak pattern, that of stomatal conductance displayed a hollow pattern, and that of transpiration rate displayed a signal-peak pattern. Water use efficiency culminated in the early morning. On the basis of two criteria of stomatal limitation of photosynthesis suggested by Farquhar and Sharkey, the predominant limiting factor of photosynthesis was the stomatal conductance of stomatal limitation in the morning (10:00–12:00). However, the midday depression of photosynthesis at noon (12:00–14:00) and the decrease in photosynthesis in the afternoon (16:00–18:00) were the results of nonstomatal limitation, such as low carboxylation capacity of the leaf mesophyll.     

Effects of water stress on Haloxylon ammodendron seedlings in the desert region of Heihe inland river watershed, Gansu Province, China

The water relation and leaf gas exchange of saxoul (Haloxylon Ammodendron Bge, a C4 shrub) seedlings were studied under water stress in 2001. Saxoul seedlings maintained high transpiration when the soil moisture was above 11%. The seedlings were able to take up water from soil with above 6 % soil water content, which was the threshold level of soil moisture for seedlings. The relationship between transpiration and potential evaporation was linear for well-watered seedlings. The decrease of soil water availability led to different degrees of down-regulation of stomatal conductance, leaf transpiration and net CO2 assimilation rate. The stomata played a relatively small part in determining the net CO2 assimilation rate for the same seedling. The relationship between net CO2 assimilation rate and transpiration was linear diurnally, and reduction scale of leaf transpiration was much bigger than that of net CO2 assimilation rate by waters tress treatments, therefore intrinsic water-use-efficiency increased. High evaporative demand increased the leaf transpiration but inhibited net CO2 assimilation rate.Because of the effect of VPD on transpiration in this region, the transpiration of well-watered and mild water stress seedlings becomes responsive to change in stomatal conductance over a wider range.     

Stomatal responses to drought at a Mediterranean site: a comparative study of co-occurring woody species differing in leaf longevity

We studied stomatal responses to decreasing predawn water potential (Psipd) and increasing leaf-to-air water vapor pressure difference (VPD) of co-occurring woody Mediterranean species with contrasting leaf habits and growth form. The species included two evergreen oaks (Quercus ilex subsp. ballota (Desf.) Samp. and Q. suber L.), two deciduous oaks (Q. faginea Lam. and Q. pyrenaica Willd.) and two deciduous shrubs (Pyrus bourgaeana Decne. and Crataegus monogyna Jacq.). Our main objective was to determine if stomatal sensitivity is related to differences in leaf life span and leaf habit. The deciduous shrubs had the least conservative water-use characteristics, with relatively high stomatal conductance and low stomatal sensitivity to soil and atmospheric drought. As a result, Psipd decreased greatly in both species during the growing season, resulting in early leaf abscission in the summer. The deciduous oaks showed intermediate water-use characteristics, having maximum stomatal conductances and CO2 assimilation rates similar to or even higher than those of the deciduous shrubs. However, they had greater stomatal sensitivity to soil drying and showed less negative Psipd values than the deciduous shrubs. The evergreen oaks, and especially the species with the greatest leaf longevity, Q. ilex, exhibited the most conservative water-use behavior, having lower maximum stomatal conductances and greater sensitivity to VPD than the deciduous species. As a result, Psipd decreased less during the growing season in the evergreens than in the deciduous species, which may contribute to greater leaf longevity by avoiding irreversible damage during the summer drought. However, the combination of low maximum CO2 assimilation rates and high stomatal sensitivity to drought must have a negative impact on the final carbon budget of leaves with a long life span.     

4个兔眼蓝莓品种光合特性对季节变化的响应

为给兔眼蓝莓品种筛选提供参考,选取‘粉蓝’、‘梯芙蓝’、‘灿烂’、‘芭尔德温’4个品种作为研究对象,在不同季节对其光合指标和叶绿素含量进行研究。结果表明:‘芭尔德温’叶片中叶绿素含量最高,‘梯芙蓝’最低。4个兔眼蓝莓品种的净光合速率、气孔导度和蒸腾速率在5—7月较高,11月最低;胞间CO2浓度和水分利用效率在11月最高,5—7月较低。‘芭尔德温’的平均净光合速率最高,‘灿烂’最低;‘灿烂’的平均气孔导度和胞间CO2浓度最大;‘粉蓝’和‘芭尔德温’的平均蒸腾速率低于‘梯芙蓝’和‘灿烂’;‘粉蓝’的平均水分利用效率最高,其次是‘芭尔德温’,‘梯芙蓝’最低。4个品种的净光合速率与蒸腾速率、气孔导度显著正相关,与胞间CO2浓度、水分利用效率显著负相关;光合有效辐射、大气温度与净光合速率、蒸腾速率及气孔导度显著正相关,与胞间CO2浓度、水分利用效率显著负相关。综上,4个品种光合特性差异显著,在不同季节其规律性特征不一致。‘芭尔德温’的净光合速率受季节影响较小,且有较大生长量;‘粉蓝’和‘芭尔德温’具有较好的节水性和耐旱性,并且在同化作用下积累干物质能力比‘梯芙蓝’和‘灿烂’强;兔眼蓝莓在5—7月具有较强的光合性能,光合有效辐射和大气温度是影响兔眼蓝莓光合作用的主要生态因子。     

Responses to water stress in two Eucalyptus globulus clones differing in drought tolerance

We evaluated drought resistance mechanisms in a drought-tolerant clone (CN5) and a drought-sensitive clone (ST51) of Eucalyptus globulus Labill. based on the responses to drought of some physiological, biophysical and morphological characteristics of container-grown plants, with particular emphasis on root growth and hydraulic properties. Water loss in excess of that supplied to the containers led to a general decrease in growth and significant reductions in leaf area ratio, specific leaf area and leaf-to-root area ratio. Root hydraulic conductance and leaf-specific hydraulic conductance decreased as water stress became more severe. During the experiment, the drought-resistant CN5 clone maintained higher leaf water status (higher predawn and midday leaf water potentials), sustained a higher growth rate (new leaf area expansion and root growth) and displayed greater carbon allocation to the root system and lower leaf-to-root area ratio than the drought-sensitive ST51 clone. Clone CN5 possessed higher stomatal conductances at moderate stress as well as higher hydraulic conductances than Clone ST51. Differences in the response to drought in root biomass, coupled with changes in hydraulic properties, accounted for the clonal differences in drought tolerance, allowing Clone CN5 to balance transpiration and water absorption during drought treatment and thereby prolong the period of active carbon assimilation.     

Leaf morphological and physiological adjustments to the spectrally selective shade imposed by anthocyanins in Prunus cerasifera

Prunus domestica L. has green leaves, whereas Prunus cerasifera Ehrh. var. atropurpurea has red leaves due to the presence of mesophyll anthocyanins. We compared morphological and photosynthetic characteristics of leaves of these species, which were sampled from shoots grafted in pairs on P. domestica rootstocks, each pair comprising one shoot of each species. Two hypotheses were tested: (1) anthocyanins protect red leaves against photoinhibition; and (2) red leaves display shade characteristics because of light attenuation by anthocyanins. Parameters were measured seasonally, during a period of increasing water stress, which caused a similar drop in shoot water potential in each species. As judged by predawn measurements of maximum PSII yield, chronic photoinhibition did not develop in either species and, despite the anthocyanic screen, the red leaves of P. cerasifera displayed lower light-adapted PSII yields and higher non-photochemical quenching than the green leaves of P. domestica. Thus, it appears that, in this system, anthocyanins afford little photoprotection. As predicted by the shade acclimation hypothesis, red leaves were thinner and had a lower stomatal frequency, area- based CO2 assimilation rate, apparent carboxylation efficiency and chlorophyll a:b ratio than green leaves. However, red leaves were similar to green leaves in conductivity to water vapor diffusion, dry-mass-based chlorophyll concentrations and carotenoid:chlorophyll ratios. The data for red leaves indicate adaptations to a green-depleted, red-enriched shade, rather than a neutral or canopy-like shade. Thus, green light attenuation by anthocyanins may impose a limitation on leaf thickness. Moreover, the selective depletion of light at wavelengths that are preferentially absorbed by PSII and chlorophyll b may lead to adjustments in chlorophyll pigment ratios to compensate for the uneven spectral distribution of internal light. The apparent photosynthetic cost associated with lost photons and reduced leaf thickness, and the absence of a photoprotective advantage, suggest that there are other, yet to be identified, benefits for permanently anthocyanic leaves of P. cerasifera.     

Photosynthesis and water-use efficiency of sugar maple (Acer saccharum) in relation to pear thrips defoliation

An experimental introduction of pear thrips (Taeniothrips inconsequens Uzel), a major defoliator in sugar maple (Acer saccharum Marsh.) forests in northeastern North America, was conducted in a field plantation to determine if compensatory gas exchange occurs in response to feeding damage by this piercing-sucking insect. Sugar maple trees were enclosed in netting (167 micro m mesh) and pear thrips adults were introduced before leaf expansion in the spring. Pear thrips reduced whole-tree leaf area by approximately 23% and reduced leaf size (both mass and area) by 20% in the upper crown. Measurements of net CO(2) assimilation rate (A(net)) and stomatal conductance (g(s)) were made on tagged foliage that was later analyzed for stable carbon isotope composition (delta(13)C) to provide estimates of short- and long-term leaf water use efficiency (WUE). Pear thrips feeding reduced A(net) for fully expanded leaves by approximately 20%, although leaf chlorophyll content and leaf mass per unit area were apparently not affected. Comparison of A(net), g(s), instantaneous WUE and leaf delta(13)C between damaged and control trees as well as visibly undamaged versus moderately damaged foliage on pear thrips-infested trees indicated that there were no effects of pear thrips feeding damage on WUE or leaf delta(13)C. Long-term WUE among sugar maple trees in the field plantation, indicated by leaf delta(13)C analysis, was related to shorter-term estimates of leaf gas exchange behavior such as g(s) and calculated leaf intercellular CO(2) concentration (C(i)). We conclude that pear thrips feeding has no effect on leaf WUE, but at the defoliation levels in our experiment, it may reduce leaf A(net), as a result of direct tissue damage or through reduced g(s). Therefore, even small reductions in leaf A(net) by pear thrips feeding damage may have an important effect on the seasonal carbon balance of sugar maple when integrated over the entire growing season.