Responses of gas exchange to reversible changes in whole-plant transpiration rate in two conifer species

This study examined the autonomy of branches with respect to the control of transpiration (E) in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata Donn) seedlings. Experiments were conducted on whole seedlings in a gas exchange system with a dual-cuvette that permitted independent manipulation and measurement of E in the upper and lower cuvettes. The value of E in one cuvette was manipulated by varying vapor pressure deficit (D) between 2.2 and 0.2 kPa, whereas D in the other cuvette was held at 2.2 kPa. Reducing D, while increasing stomatal conductance (gs), resulted in an overall decrease in E. In western red cedar, this decrease was almost threefold, and in Douglas-fir, approximately fourfold. In well-watered western red cedar, a reduction of whole-plant E by 46% (brought about by reducing D in the upper cuvette) resulted in a 12% increase in gs, a 12% increase in E and a 7% increase in net assimilation (A) of untreated foliage in the lower cuvette. Responses of gs, E and A of untreated foliage were similar irrespective of whether foliage was at the top or bottom of the seedling. When D in the treatment cuvette was restored to 2.2 kPa, gs, E and A of foliage in the untreated cuvette returned to pretreatment values. In contrast, in well-watered Douglas-fir, there was almost no change in gs, E or A of untreated foliage in one cuvette when D in the other cuvette was reduced, causing a 52% reduction in whole-plant E. However, similar manipulations on drought-stressed Douglas-fir led to 7-19% increases in gs, E and A of untreated foliage. In well-watered western red cedar, daytime leaf water potential (Psil) was maintained near -0.9 MPa over a wide range of D, whereas Psil of Douglas-fir decreased from -1.2 to -1.5 MPa as D increased. The tighter (isohydric) regulation of Psil in western red cedar may partly explain its greater stomatal response to D and variation in whole-plant E compared with Douglas-fir. In response to a reduction in E, measured increases in Psil and gs of unmanipulated foliage were less than predicted by a model assuming complete hydraulic connectivity of foliage. Our results suggest the foliage of both species is partially autonomous with respect to water.     

Photosynthesis and light-use efficiency by plants in a Canadian boreal forest ecosystem

Measurements of the photosynthetic response to midsummer irradiance were made for 11 species representing the dominant trees, understory shrubs, herbaceous plants and moss species in an old black spruce (Picea mariana (Mill.) B.S.P.) boreal forest ecosystem. Maximum rates of photosynthesis per unit foliage area at saturating irradiance, A(max), were highest for aspen (Populus tremuloides Michx.), reaching 16 micromol m(-2) s(-1). For tamarack (Larix laricina (Du Roi) K. Kock) and P. mariana, Amax was only 2.6 and 1.8 micromol m(-2) s(-1), respectively. Values of A(max) for understory shrubs and herbaceous plants were clustered between 9 and 11 micromol m(-2) s(-1), whereas A(max) of feather moss (Pleurozium schreberi (Brid.) Mitt.) reached only 1.9 micromol m(-2) s(-1). No corrections were made for differences in shoot structure, but values of photosynthetic light-use efficiency were similar for most species (70-80 mmol CO2 mol(-1)); however, they were much lower for L. laricina and P. mariana (15 mmol CO2 mol(-1)) and much higher for P. schreberi (102 m;mol CO2 mol(-1)). There was a linear relationship between Amax and foliage nitrogen concentration on an area basis for the broad-leaved species in the canopy and understory, but the data for P. mariana, L. laricina and P. schreberi fell well below this line. We conclude that it is not possible to scale photosynthesis from leaves to the canopy in this ecosystem based on a single relationship between photosynthetic rate and foliage nitrogen concentration.     

Short-term effects of fertilization on photosynthesis and leaf morphology of field-grown loblolly pine following long-term exposure to elevated CO(2) concentration

We examined effects of a first nitrogen (N) fertilizer application on upper-canopy needle morphology and gas exchange in approximately 20-m-tall loblolly pine (Pinus taeda L.) exposed to elevated carbon dioxide concentration ([CO(2)]) for 9 years. Duke Forest free-air CO(2) enrichment (FACE) plots were split and half of each ring fertilized with 112 kg ha(-1) elemental N applied in two applications in March and April 2005. Measurements of needle length (L), mass per unit area (LMA), N concentration (N(l)) on a mass and an area basis, light-saturated net photosynthesis per unit leaf area (A(a)) and per unit mass (A(m)), and leaf conductance (g(L)) began after the second fertilizer application in existing 1-year-old foliage (F(O)) and later in developing current-year first-flush (F(C1)) and current-year second-flush (F(C2)) foliage. Elevated [CO(2)] increased A(a) by 43 and 52% in F(O) and F(C1) foliage, respectively, but generally had no significant effect on any other parameter. Fertilization had little or no significant effect on L, LMA, A or g(L) in F(O) foliage; although N(l) was significantly higher in fertilized trees by midsummer. In contrast, fertilization resulted in large increases in L, N(l), and A in F(C1) and F(C2) foliage, increasing A(a) by about 20%. These results suggest that, although both needle age classes accumulate N following fertilization, they use it differently-current-year foliage incorporates N into photosynthetic machinery, whereas 1-year-old foliage serves as an N store. There were no significant interaction effects of elevated [CO(2)] and fertilization on A. Elevated [CO(2)] increased the intercept of the A:N(l) relationship but did not significantly affect the slope of the relationship in either foliage age class.     

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.     

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.     

Effects of carbon dioxide, fertilization, and irrigation on photosynthetic capacity of loblolly pine trees

Branches of nine-year-old loblolly pine trees grown in a 2 x 2 factorial combination of fertilization and irrigation were exposed for 11 months to ambient, ambient + 175, or ambient + 350 micro mol mol(-1) CO(2). Rates of light-saturated net photosynthesis (A(max)), maximum stomatal conductance to water vapor (g(max)), and foliar nitrogen concentration (% dry mass) were assessed monthly from April 1993 until September 1993 on 1992 foliage (one-year-old) and from July 1993 to March 1994 on 1993 foliage (current-year). Rates of A(max) of foliage in the ambient + 175 CO(2) treatment and ambient + 350 were 32-47 and 83-91% greater, respectively, than that of foliage in the ambient CO(2) treatment. There was a statistically significant interaction between CO(2) treatment and fertilization or irrigation treatment on A(max) on only one measurement date for each age class of foliage. Light-saturated stomatal conductance to water vapor (g(max)) was significantly affected by CO(2) treatment on only four measurement dates. Light-saturated g(max) in winter was only 42% of summer g(max) even though soil water during winter was near field capacity and evaporative demand was low. Fertilization increased foliar N concentration by 30% over the study period when averaged across CO(2) treatments. During the study period, the ambient + 350 CO(2) treatment decreased average foliar N concentration of one-year-old foliage in the control, irrigated, fertilized and irrigated + fertilized plots by 5, 6.4, 9.6 and 11%, respectively, compared with one-year-old foliage in the corresponding ambient CO(2) treatments. The percent increase in A(max) due to CO(2) enrichment was similar in all irrigation and fertilization treatments and the effect persisted throughout the 11-month study period for both one-year-old and current-year foliage.     

Photosynthetic capacity and nitrogen partitioning in foliage of the evergreen shrub Daphniphyllum humile along a natural light gradient

We examined the effects of leaf age and mutual shading on the morphology, photosynthetic properties and nitrogen (N) allocation of foliage of an evergreen understory shrub, Daphniphyllum humile Maxim, growing along a natural light gradient in a deciduous Fagus crenata-dominated forest in Japan. Seedlings in high-light environments were subject to greater mutual shading and 1-year-old foliage survival was lower than in seedlings in low-light environments, indicating that the survival rates of foliage were related to the degree of mutual shading. Although specific leaf area (SLA) in current- and 1-year-old foliage was curvilinearly related to daily photosynthetic photon flux (PPF), SLA was unaffected by leaf age, indicating that foliage in D. humile may not acclimate morphologically to annual changes in light caused by mutual shading. Light-saturated net photosynthetic rates (Pmax) were correlated with daily PPF in current-year foliage. In addition, a strong, positive relationship was found between nitrogen concentration per unit leaf area and Pmax. In contrast, the relationship among PPF, N and photosynthetic parameters in 1-year old foliage was weak because of the strong remobilization of N from older leaves to current-year foliage in plants growing in high light. However, the relationship between daily PPF and both photosynthetic N-use efficiency and the ratio of maximum electron transport rate to maximum carboxylation rate did not differ between current-year and 1-year-old foliage, suggesting that these responses help maintain a high photosynthetic efficiency even in older foliage. We conclude that D. humile maximizes whole-plant carbon gain by maintaining a balance among photosynthetic functions across wide ranges of leaf ages and light environments.     

遮光对野鸦椿幼苗生长和光合生理特性的影响

为了解不同光照条件下野鸦椿幼苗的生长发育状况和光合生理特性,探讨野鸦椿幼苗生长发育的最佳光照条件.采用遮光方式模拟不同光照强度(全光照,遮光30％、60％和90％),研究野鸦椿幼苗的生长、生物量、光合色素含量、生理指标和光合日变化等光合特性的响应规律.结果表明:随着遮光程度的增加,野鸦椿幼苗的生长量均高于对照,以遮光6...     

Thinning,fertilization, and crown position interact to control physiological responses of loblolly pine

To examine physiological responses to thinning, fertilization, and crown position, we measured net photosynthesis (P(n)), transpiration (E), vapor pressure difference (VPD), stomatal conductance (g(s)), and xylem pressure potential (Psi(1)) between 0930 and 1130 h under ambient conditions in the upper and lower crowns of a 13-year-old loblolly pine (Pinus taeda L.) plantation six years (1994) after the treatments were applied. Photosynthetic photon flux density (PPFD) and air temperature (T(a)) within the canopy were also recorded. Needle P(n) of thinned trees was significantly enhanced by 22-54% in the lower crown, because canopy PPFD increased by 28-52%. Lower crown foliage of thinned plots also had higher E and g(s) than foliage of unthinned plots, but thinning had no effect on needle Psi(1) and predawn xylem pressure potential (0430-0530 h; Psi(pd)). Tree water status did not limit P(n), E and g(s) during the late-morning measurements. Fertilization significantly decreased within-canopy PPFD and T(a). Needle Psi(1) was increased in fertilized stands, whereas P(n), E and g(s) were not significantly altered. Upper crown foliage had significantly greater PPFD, P(n), VPD, g(s), E, and more negative Psi(1) than lower crown foliage. In both crown positions, needle P(n) was closely related to g(s), PPFD and T(a) (R(2) = 0.77 for the upper crown and 0.82 for the lower crown). We conclude that (1) silvicultural manipulation causes microclimate changes within the crowns of large trees, and (2) needle physiology adjusts to the within-crown environmental conditions.     

Photosynthetic responses of cottonwood seedlings grown in glacial through future atmospheric [CO2] vary with phosphorus supply

Plants often exhibit proportionately larger photosynthetic responses to the transition from glacial to modern [CO(2)] than from modern to future [CO(2)]. Although this pattern may reflect increased nutrient demand with increasing [CO(2)], few studies have examined the role of nutrient supply in regulating responses to the range of [CO(2)] from glacial to future [CO(2)]. In this study, we examined the effects of P supply (0.004-0.5 mM) on photosynthetic responses of Populus deltoides (cottonwood) seedlings to glacial (200 micromol mol(-1)), modern (350 μmol mol(-1)) and future (700 micromol mol(-1)) [CO(2)]. The A(sat) (light-saturated net photosynthetic rates at the growth [CO(2)]) response to future [CO(2)] decreased with decreasing P supply such that there was no response at the lowest P supply. However, P supply did not affect A(sat) responses to an increase from glacial to modern [CO(2)]. Photosynthetic capacity [e.g., final rubisco activity, apparent, maximal Rubisco-limited rate of photosynthesis (V(cmax)), apparent, maximal electron transport-limited rate of photosynthesis (J(max))], stomatal conductance (g(s)) and leaf P generally increased with increasing P supply but decreased with increasing [CO(2)]. Measures of carbohydrate sink capacity (e.g., leaf mass per unit leaf area, leaf starch) increased with both increasing P supply and increasing [CO(2)]. Changes in V(cmax) and g(s) together accounted for 78% of the variation in A(sat) among [CO(2)] and P treatments, suggesting significant biochemical and stomatal controls on photosynthesis. However, A(sat) responses to increasing [CO(2)] did not reflect the changes in the carbohydrate sink capacity. These results have important implications because low P already constrains responses to increasing [CO(2)] in many ecosystems, and our results suggest that the P demand will increasingly affect A(sat) in cottonwood as [CO(2)] continues to increase.     

Partial shoot removal increases net CO(2) assimilation and alters water relations of Citrus seedlings

Effects of defoliation on partial shoot removal by decapitation on seedling growth, water use and net gas exchange of remaining basal leaves, were examined in Citrus spp. Shoot and root growth rates were manipulated to test for effects of growth demands on net gas exchange. Partially defoliated plants had higher leaf pressure potentials, root conductivities and rates of water use than intact control plants. Shoot regrowth occurred at the expense of root loss. Basal leaves on defoliated plants consistently had higher rates of CO(2) assimilation (A) than leaves on intact plants. Stomatal conductance (g(s)) changed little after defoliation so the higher A of leaves on defoliated plants lowered the ratio of intercellular to ambient CO(2) concentration (C(i)/C(a)) in the mesophyll. In some cases, g(s) increased with A in defoliated plants but C(i)/C(a) was not affected. Stomatal conductance only limited A when intact seedlings were stressed by root confinement in small pots or when leaves were exposed to high vapor pressure deficits during gas exchange measurements. Increased carbon demand for shoot regrowth increased photosynthetic capacity and was more important than stomatal responses in determining A after partial shoot loss.     

Temperature effects on wood anatomy, wood density, photosynthesis and biomass partitioning of Eucalyptus grandis seedlings

Wood density, a gross measure of wood mass relative to wood volume, is important in our understanding of stem volume growth, carbon sequestration and leaf water supply. Disproportionate changes in the ratio of wood mass to volume may occur at the level of the whole stem or the individual cell. In general, there is a positive relationship between temperature and wood density of eucalypts, although this relationship has broken down in recent years with wood density decreasing as global temperatures have risen. To determine the anatomical causes of the effects of temperature on wood density, Eucalyptus grandis W. Hill ex Maiden seedlings were grown in controlled-environment cabinets at constant temperatures from 10 to 35 degrees C. The 20% increase in wood density of E. grandis seedlings grown at the higher temperatures was variously related to a 40% reduction in lumen area of xylem vessels, a 10% reduction in the lumen area of fiber cells and a 10% increase in fiber cell wall thickness. The changes in cell wall characteristics could be considered analogous to changes in carbon supply. Lumen area of fiber cells declined because of reduced fiber cell expansion and increased fiber cell wall thickening. Fiber cell wall thickness was positively related to canopy CO2 assimilation rate (Ac), which increased 26-fold because of a 24-fold increase in leaf area and a doubling in leaf CO2 assimilation rate from minima at 10 and 35 degrees C to maxima at 25 and 30 degrees C. Increased Ac increased seedling volume, biomass and wood density; but increased wood density was also related to a shift in partitioning of seedling biomass from roots to stems as temperature increased.     

Developmental decline in height growth in Douglas-fir

The characteristic decline in height growth that occurs over a tree's lifespan is often called "age-related decline." But is the reduction in height growth in aging trees a function of age or of size? We grafted shoot tips across different ages and sizes of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees to determine whether the decline in height growth is mediated by tree size or by the age of the apical meristem. We also evaluated whether reduced carbon assimilation plays an important role in height growth decline. In one experiment we cut shoot tips from old-growth, young-mature and seedling trees and grafted them onto 2-year-old graft-compatible rootstock in a seed orchard in Lebanon, Oregon. In another experiment we performed reciprocal grafts between lateral branches of old-growth trees accessible from the canopy crane at Wind River, Washington and young-mature trees in a nearby plantation. We measured growth (diameter and elongation of the dominant new stem) and mortality annually for three years in the Seed Orchard experiment and for two years in the Reciprocal Graft experiment. In the Seed Orchard experiment we also measured photosynthetic capacity (determined from the response of net carbon assimilation to the intercellular CO(2) concentration of the leaf, or A/C(i) curves), leaf mass per area (LMA) and carbon isotope composition (delta(13)C) of cellulose in 1-year-old foliage. Grafting caused changes in both growth and physiology of the grafted stems. Within two years after grafting, growth and physiology of all combinations of scions and rootstock exhibited characteristics of the rootstock. In some cases, the change in growth was dramatic-cuttings from old-growth trees showed a 10-fold increase in stem elongation rate within 2 years of grafting onto seedling rootstock. Similarly, carbon isotope discrimination of new foliage on shoots from old-growth trees increased by nearly 3 per thousand and 2 per thousand after grafting onto young-mature and seedling rootstock, respectively, whereas discrimination decreased by a similar magnitude in scions from young-mature trees after grafting on old- growth trees. Furthermore, differences in carbon assimilation estimated from carbon isotope discrimination and A/C(i )relationships were small relative to growth differences. Our results confirm that size, not age, drives developmental changes in height growth in Douglas-fir. Reduced carbon assimilation does not play an important role in height growth decline.     

Effects of elevated CO(2) concentration and nutrition on net photosynthesis,stomatal conductance and needle respiration of field-grown Norway spruce trees

To study the effects of elevated CO(2) on gas exchange, nonstructural carbohydrate and nutrient concentrations in current-year foliage of 30-year-old Norway spruce (Picea abies (L.) Karst.) trees, branches were enclosed in ventilated, transparent plastic bags and flushed with ambient air (mean 370 &mgr;mol CO(2) mol(-1); control) or ambient air + 340 &mgr;mol CO(2) mol(-1) (elevated CO(2)) during two growing seasons. One branch bag was installed on each of 24 selected trees from control and fertilized plots. To reduce the effect of variation among trees, results from each treated branch were compared with those from a control branch on the same whorl of the same tree. Elevated CO(2) increased rates of light-saturated photosynthesis on average by 55% when measured at the treatment CO(2) concentration. The increase was larger in shoots with high needle nitrogen concentrations than in shoots with low needle nitrogen concentrations. However, shoots grown in elevated CO(2) showed a decrease in photosynthetic capacity compared with shoots grown in ambient CO(2). When measured at the internal CO(2) concentration of 200 &mgr;mol CO(2) mol(-1), photosynthetic rates of branches in the elevated CO(2) treatments were reduced by 8 to 32%. The elevated CO(2) treatment caused a 9 to 20% reduction in carboxylation efficiency and an 18% increase in respiration rates. In response to elevated CO(2), starch, fructose and glucose concentrations in the needles increased on average 33%, whereas concentrations of potassium, nitrogen, phosphorus, magnesium and boron decreased. Needle nitrogen concentrations explained 50-60% of the variation in photosynthesis and CO(2) acclimation was greater at low nitrogen concentrations than at high nitrogen concentrations. We conclude that the enhanced photosynthetic rates found in shoots exposed to elevated CO(2) increased carbohydrate concentrations, which may have a negative feedback on the photosynthetic apparatus and stimulate cyanide-resistant respiration. We also infer that the decrease in nutrient concentrations of needles exposed to elevated CO(2) was the result of retranslocation of nutrients to other parts of the branch or tree.     

Photosynthetic responses of field-grown Pinus radiata trees to artificial and aphid-induced defoliation

The phloem-feeding aphid Essigella californica represents a potential threat to the productivity of Pinus radiata plantations in south-eastern Australia. Five- and nine-year-old field trials were used to characterize the effects of artificial and natural aphid-induced (E. californica) defoliation, respectively, on shoot photosynthesis and growth. Photosynthetic capacity (A(max)) was significantly greater following a 25% (D25) (13.8 μmol m(-2) s(-1)) and a 50% (D50) (15.9 μmol m(-2) s(-1)) single-event upper-crown artificial defoliation, 3 weeks after defoliation than in undefoliated control trees (12.9 μmol m(-2) s(-1)). This response was consistently observed for up to 11 weeks after the defoliation event; by Week 16, there was no difference in A(max) between control and defoliated trees. In the D50 treatment, this increased A(max) was not sufficient to fully compensate for the foliage loss as evidenced by the reduced diameter increment (by 15%) in defoliated trees 36 weeks after defoliation. In contrast, diameter increment of trees in the D25 treatment was unaffected by defoliation. The A(max) of trees experiencing upper-crown defoliation by natural and repeated E. californica infestations varied, depending on host genotype. Despite clear differences in defoliation levels between resistant and susceptible genotypes (17 vs. 35% of tree crown defoliated, respectively), growth of susceptible genotypes was not significantly different from that of resistant genotypes. The observed increases in A(max) in the lower crown of the canopy following attack suggested that susceptible genotypes were able to partly compensate for the loss of foliage by compensatory photosynthesis. The capacity of P. radiata to regulate photosynthesis in response to natural aphid-induced defoliation provides evidence that the impact of E. californica attack on stem growth will be less than expected, at least for up to 35% defoliation.     

遮荫对东北红豆杉苗期生长及光合特性的影响

在100%、70%、40%和10%4种光照条件下,对东北红豆杉苗期生长及光合特性进行了研究。结果表明:在100%、70%光照下,净光合速率(Pn)日变化呈现"双峰型"曲线,有明显的"光午休"现象,后者较前者的"光午休"作用减弱了17.7个百分点;强光、高温、低湿是引起"光午休"的主要环境因子。在70%光照下,苗高、地径的生长量最大,净光合速率和水分利用率最高。70%光照是东北红豆杉苗期生长的适宜光环境。     

Changes in rates of photosynthesis and respiration during needle development of loblolly pine

Net photosynthetic rates of developing foliage and one-year-old foliage of loblolly pine (Pinus taeda L.) were measured under field conditions. In the subsequent year, net photosynthesis and dark respiration rates of current-year and one-year-old foliage were measured under controlled environmental conditions. Loblolly pine foliage grows slowly, reaching its final size 3.5 to 4 months after bud burst. Positive rates of net photosynthesis were recorded when the foliage was 13 and 18% of final length, in the controlled-environment and field study, respectively. However, because of high rates of dark respiration during the initial growth period, a positive diurnal carbon balance did not occur until foliage was about a third of final length (40 days after bud burst). Two months after bud burst, when foliage was about 55% of final length, its photosynthetic capacity exceeded that of one-year-old foliage. The highest rates of net photosynthesis were achieved when foliage was more than 90% fully expanded.     

Stomatal conductance alone does not explain the decline in foliar photosynthetic rates with increasing tree age and size in Picea abies and Pinus sylvestris

Foliar light-saturated net assimilation rates (A) generally decrease with increasing tree height (H) and tree age (Y), but it is unclear whether the decline in A is attributable to size- and age-related modifications in foliage morphology (needle dry mass per unit projected area; M(A)), nitrogen concentration, stomatal conductance to water vapor (G), or biochemical foliage potentials for photosynthesis (maximum carboxylase activity of Rubisco; V(cmax)). I studied the influences of H and Y on foliage structure and function in a data set consisting of 114 published studies reporting observations on more than 200 specimens of various height and age of Picea abies (L.) Karst. and Pinus sylvestris L. In this data set, foliar nitrogen concentrations were independent of H and Y, but net assimilation rates per unit needle dry mass (A(M)) decreased strongly with increasing H and Y. Although M(A) scaled positively with H and Y, net assimilation rates per unit area (A(A) = M(A) x A(M)) were strongly and negatively related to H, indicating that the structural adjustment of needles did not compensate for the decline in mass-based needle photosynthetic rates. A relevant determinant of tree height- and age-dependent modifications of A was the decrease in G. This led to lower needle intercellular CO2 concentrations and thereby to lower efficiency with which the biochemical photosynthetic apparatus functioned. However, V(cmax) per unit needle dry mass and area strongly decreased with increasing H, indicating that foliar photosynthetic potentials were lower in larger trees at a common intercellular CO2 concentration. Given the constancy of foliar nitrogen concentrations, but the large decline in apparent V(cmax) with tree size and age, I hypothesize that the decline in Vcmax results from increasing diffusive resistances between the needle intercellular air space and carboxylation sites in chloroplasts. Increased diffusive limitations may be the inevitable consequence of morphological adaptation (changes in M(A) and needle density) to greater water stress in needles of larger trees. Foliage structural and physiological variables were nonlinearly related to H and Y, possibly because of hyperbolic decreases in shoot hydraulic conductances with increasing tree height and age. Although H and Y were correlated, foliar characteristics were generally more strongly related to H than to Y, suggesting that increases in height rather than age are responsible for declines in foliar net assimilation capacities.     

Modeling the effect of physiological responses to green pruning on net biomass production of Eucalyptus nitens

Green pruning of Eucalyptus nitens (Deane and Maiden) Maiden increases instantaneous rates of light-saturated CO(2) assimilation (A), and changes patterns of total leaf area and foliage distribution. We investigated the importance of such changes on the rate of recovery of growth following pruning. A simple process-based model was developed to estimate daily net biomass production (G(d)) of three-year-old plantation-grown trees over a 20-month period. The trees had been pruned by removal of 0, 50 or 70% of the length of green crown, equivalent to removal of 0, 55 or 88% of leaf area, respectively, when the plantation verged on canopy closure. Total G(d) was reduced by only 20% immediately following the 50%-pruning treatment, as a result of both the high leaf dark respiration and low A in the portion of the crown removed compared to the top of the crown. Pruning at the time of canopy closure preempted a natural and rapid decline in G(d) of the lower crown. Although leaf area index (L) was approximately 6.0 at the time of pruning, high light interception (95%) occurred with an L of 4.0. The 50%-pruning treatment reduced L to 3.5, but the physiological responses to pruning were sufficient to compensate fully for the reduction in intercepted radiation within 110 days of pruning. The 70%-pruning treatment reduced L to 1.9, and reduced G(d) by 77%, reflecting the removal of branches with high A in the mid and upper crown. Physiological responses to the 70%-pruning treatment were insufficient to increase G(d) to the value of unpruned trees during the study. Model sensitivity analysis showed that increases in A following pruning increased G(d) by 20 and 25% in the 50- and 70%-pruned trees, respectively, 20 months after pruning. Changes in leaf area/foliage distribution had a greater effect on G(d) of 50%-pruned trees (47% increase) than did changes in A. However, the reduction in photosynthetic potential associated with the 70%-pruning treatment resulted in only small changes in leaf area/foliage distribution, which consequently had little effect on G(d). The effects of physiological processes occurring within the crown and in response to green pruning on G(d) are discussed with respect to pruning of plantations.     

Using combined measurements for comparison of light induction of stomatal conductance, electron transport rate and CO2 fixation in woody and fern species adapted to different light regimes

We aimed to understand the relation of photosynthetic rate (A) with g(s) and electron transport rate (ETR) in species of great taxonomic range and light adaptation capability during photosynthetic light induction. We studied three woody species (Alnus formosana, Ardisia crenata and Ardisia cornudentata) and four fern species (Pyrrosia lingus, Asplenium antiquum, Diplazium donianum and Archangiopteris somai) with different light adaptation capabilities. Pot-grown materials received 100 and/or 10% sunlight according to their light adaptation capabilities. At least 4 months after light acclimation, CO(2) and H(2)O exchange and chlorophyll fluorescence were measured simultaneously by equipment in the laboratory. In plants adapted or acclimated to low light, dark-adapted leaves exposed to 500 or 2000 μmol m(-2) s(-1) photosynthetic photon flux (PPF) for 30 min showed low gross photosynthetic rate (P(g)) and short time required to reach 90% of maximum P(g) (). At the initiation of illumination, two broad-leaved understory shrubs and the four ferns, especially ferns adapted to heavy shade, showed higher stomatal conductance (g(s)) than pioneer tree species; materials with higher g(s) had short at both 500 and 2000 μmol m(-2) s(-1) PPF. With 500 or 2000 μmol m(-2) s(-1) PPF, the g(s) for the three woody species increased from 2 to 30 min after the start of illumination, but little change in the g(s) of the four ferns. Thus, P(g) and g(s) were not correlated for all material measured at the same PPF and induction time. However, P(g) was positively correlated with ETR, even though CO(2) assimilation may be influenced by stomatal, biochemical and photoinhibitory limitations. In addition, was closely related to time required to reach 90% maximal ETR for all materials and with two levels of PPF combined. Thus, ETR is a good indicator for estimating the light induction of photosynthetic rate of species, across a wide taxonomic range and light adaptation and acclimation capability.