Abscisic acid accumulation in leaves of two contrasting hybrid poplar clones affected by nitrogen fertilization plus cyclic flooding and soil drying

Cuttings of hybrid Populus clones Tristis and Eugenei growing in pots in a greenhouse were treated with nitrogen fertilizer at two rates and subjected to repeated soil flooding or drying. Periodically, gas exchange measurements and radioimmunoassays, to determine abscisic acid (ABA) concentrations, were made on recently mature leaves.In both clones, photosynthesis and stomatal conductance were depressed five days after flooding, but leaf ABA concentrations remained relatively constant. In contrast, an initial, 9-day period of soil drying resulted in substantial ABA accumulation in leaves, which closely correlated with declines in photosynthesis and conductance. A second soil drying cycle of up to 9 days was less effective in modifying gas exchange and leaf ABA concentrations. High-N supply stimulated leaf ABA production as the soil dried. On the resumption of watering, gas exchange in Tristis recovered fully and rapidly and leaf ABA concentrations quickly returned to control values, whereas gas exchange in Eugenei recovered slowly and leaf ABA concentrations remained high for longer.Gas exchange in Eugenei was unaffected by soil drying until leaf ABA concentrations exceeded 100 ng g(dw) (-1), whereas Tristis showed a reduction in stomatal conductance and photosynthesis at leaf ABA concentrations of only 10 ng g(dw) (-1). A rise in internal CO(2) concentrations was associated with increased leaf ABA concentrations in Tristis, but not in Eugenei. Clonal differences in the relationship between gas exchange and leaf ABA concentration suggest contrasting physiological strategies for survival under prolonged drying conditions.     

Growth and development during the establishment year of two Populus clones with contrasting morphology and phenology

Weekly morphological measurements of trees in permanent growth plots and periodic destructive sampling were used to monitor growth and development of two Populus clones with contrasting morphology and phenology during the establishment year in a short-rotation, intensive-culture system. Tristis (P. tristis Fisch. x P. balsamifera L.) grew rapidly for 48 days before setting bud in July. By contrast, Eugenei (P. x euramericana (Dode) Guinier) grew at a slower rate than Tristis, but maintained this rate for 75 days before setting bud in September. By early October, the total leaf area and dry weight of Eugenei exceeded that of Tristis by 39 and 11%, respectively. In addition, Eugenei had a greater harvest index than Tristis throughout most of the growing season because a larger proportion of photosynthate produced was directed to shoot growth; however, a high shoot/root ratio in Eugenei predisposed it to water stress. Differences in aboveground biomass between clones were largely attributable to clonal differences in seasonal leaf area development.     

Differential physiological and morphological responses of two hybrid Populus clones to water stress

Hardwood cuttings of Populus clones Tristis #1 and Eugenei were grown in pots in a controlled environment chamber to observe early patterns of growth and water relations in response to changing conditions of water stress. Height and dry weight growth, dry matter partitioning, leaf area production, stomatal conductance and leaf water potential were measured periodically during the 73-day experiment. The two clones reacted in a similar way to an initial period of stress, showing reduced growth, stomatal conductance and leaf water potential. However, when stress was interrupted by ample watering and then reimposed, substantial differences between the clones were evident. Growth of Eugenei fully recovered after stress was relieved, especially leaf growth, but when water deficiency was reimposed, the plants wilted and some leaves died. Tristis #1, in contrast, showed a greater adaptation to changing stress conditions; it grew less than Eugenei after drought was interrupted, but showed little adverse effect of renewed water deficits. These responses were partially explained by the higher root/leaf weight ratios of Tristis #1 which enabled it to maintain a more favorable plant water status than Eugenei.     

Comparison of photosynthetic induction and transient limitations during the induction phase in young and mature leaves from three poplar clones

We tested the hypothesis that leaf age affects photosynthetic induction, because conductance to CO2 diffusion usually decreases with increasing leaf age. Photosynthetic inductions, primarily determined by the light modulation of Rubisco activity and stomatal opening, were investigated in both young and mature leaves, as defined by leaf plastochron index (LPI), from three poplar clones: Populus alba L., P. nigra L. and P. x euramericana (Dode) Guinier. In all clones, maximum assimilation rates (A max), maximum stomatal conductance (G Smax) and dark respiration rates (RD) were higher in young leaves (LPI = 3-5) than in mature leaves (LPI = 10-14), and A max decreased from P. alba via P. x euramericana to P. nigra. The clones with high photosynthetic capacity had low induction states 60 s after leaf illumination (IS60; indicating a slow initial induction phase), and required less time to reach 90% photosynthetic induction (T90). In contrast, the clone with the lowest photosynthetic capacity (P. nigra) exhibited high IS60 (high initial induction state) but a long induction time (high T90). A comparison of mature leaves with young leaves revealed significantly (P < 0.01) lower IS60 values in mature leaves of P. nigra only, and significantly higher T90 values in mature leaves of P. alba only. In all clones, young leaves exhibited a lower percentage of maximum transient stomatal limitation during photosynthetic induction (4-9%) compared with mature leaves (16-30%). Transient biochemical limitation, assessed on the basis of the time constants of Rubisco activation (tau), was significantly higher in mature leaves than in young leaves of P. alba; whereas there were no significant differences in tau between young and mature leaves of the other poplar clones. Thus, our hypothesis that leaf age affects photosynthetic induction was confirmed at the level of transient stomatal limitation, which was significantly higher in mature leaves than in young leaves in all clones. For the induction parameters IS60, T90 and tau, photosynthetic induction was more clone-specific and was dependent on leaf age only in some cases, an observation that may apply to other tree species.     

Effects of irrigation and coppicing on above-ground growth, physiology, and fine-root dynamics of two field-grown hybrid poplar clones

To determine the effects of irrigation and coppicing on above- and below-ground growth dynamics, a plantation of Populus × euramericana cv. ‘Eugenei’ and Populus tristis × Populus balsamifera cv. ‘Tristis 1’ was established in May 1984 on a level site in East Lansing, MI, USA. Supplemental water in the form of drip irrigation was applied to half the trees beginning the first growing season. All trees were cut down in March 1988 and the stumps allowed to coppice. Pre- and post-coppice height and diameter growth of ‘Eugenei’ always exceeded that of ‘Tristis’, and the former clone showed a positive response to irrigation, whereas the latter did not. The greater growth of ‘Eugenei’ was primarily due to its full use of the growing season. Post-coppice rates of photosynthesis were not affected by irrigation in either clone, but stomatal conductances were reduced in non-irrigated trees. Analysis of microvideo images taken in minirhizotron tubes buried in the soil close to the trees showed that ‘Tristis’ produced a greater length and number of fine roots in the top 30 cm of soil than ‘Eugenei’, regardless of treatment. Irrigated trees consistently produced more fine roots at 0–30 cm soil depth than non-irrigated trees only in ‘Eugenei’, but both clones showed greater fine-root production in non-irrigated trees at 30–100 cm. Both clones also showed substantial fine-root production in the spring immediately following coppicing, with no evidence of a shock-induced dieback of roots. The root systems of these two poplar genotypes apparently contain sufficient carbon and nitrogen reserves to fuel a spring flush of fine-root growth, even though the tops have been severed during the dormant season.     

Photosynthetic productivity of aspen clones varying in sensitivity to tropospheric ozone

Rooted cuttings from three aspen (Populus tremuloides Michx.) clones (216, 271 and 259, classified as high, intermediate and low in O(3) tolerance, respectively) were exposed to either diurnal O(3) profiles simulating those of Michigan's Lower Peninsula (episodic treatments), or diurnal square-wave O(3) treatments in open-top chambers in northern Michigan, USA. Ozone was dispensed in chambers ventilated with charcoal-filtered (CF) air. In addition, seedlings were compared to rooted cuttings in their response to episodic O(3) treatments. Early in the season, O(3) caused decreased photosynthetic rates in mature leaves of all clones, whereas only the photosynthetic rates of recently mature leaves of the O(3)-sensitive Clone 259 decreased in response to O(3) exposure. During midseason, O(3) caused decreased photosynthetic rates of both recently mature and mature leaves of the O(3)-sensitive Clone 259, but it had no effect on the photosynthetic rate of recently mature leaves of the O(3)-tolerant Clone 216. Late in the season, however, photosynthetic rates of both recently mature and mature leaves of Clone 216 were lower than those of the control plants maintained in CF air. Ozone decreased the photosynthetic rate of mature leaves of Clone 271, but it increased or had no effect on the photosynthetic rate of recently mature leaves. Photosynthetic response patterns of seedlings to O(3) treatment were similar to those of the clones, but total magnitude of the response was less, perhaps reflecting the diverse genotypes of the seedling population. Early leaf abscission was observed in all clones exposed to O(3); however, Clones 216 and 259 lost more leaf area than Clone 271. By late August, leaf area in the highest O(3) treatment had decreased relative to the controls by 26, 24 and 9% for Clones 216, 259 and 271, respectively. Ozone decreased whole-tree photosynthesis in all clones, and the decrease was consistently less in Clone 271 (23%) than in Clones 216 (56%) and 259 (56%), and was accompanied by declines in total biomass of 19, 28 and 47%, respectively. The relationship between biomass and whole-tree photosynthesis indicates that the negative impact of O(3) on biomass in the clones was determined largely by lower photosynthetic productivity of the foliage, rather than by potential changes in the carbon relations of other plant organs.     

Water stress, photosynthesis and early growth patterns of cuttings of three Populus clones

Photosynthetic attributes, leaf area and early root growth patterns were studied in three Populus clones to identify traits associated with superior growth potential on sites where water could be a limiting factor. It was found that early root growth and superior leaf area production were more closely related to growth potential than were photosynthetic capacity or carboxylation efficiency. A hybrid clone of Populus nigra var. charkowiensis (syn. P. nigra var. plantierensis) x P. nigra cv. 'Incrassata' (NE308) had more leaf area production and greater root system development in both wet and dry soil than did a P. trichocarpa clone (T6) and a P. balsamifera clone (B3). Despite greater above- and below-ground productivity, plants of clone NE308 had significantly lower photosynthetic capacity and carboxylation efficiency and a slightly higher CO(2) compensation point than plants of clones T6 and B3. Rapid early leaf and root growth appear to be key attributes associated with productivity in these clones regardless of soil water availability.     

Evaluation of sampling schemes for estimating instantaneous whole-tree photosynthesis in Populus clones: a modeling approach

We evaluated several sampling schemes for estimating instantaneous whole-tree photosynthesis of 1-year-old Populus clones. Growth of two clones was simulated under varying weather conditions and leaf orientation scenarios providing photosynthetic data on a leaf-by-leaf basis throughout the growing season. Simple random sampling, stratified random sampling and a series of physiologically based sampling schemes were evaluated using either whole-leaf photosynthesis or photosynthetic rate (i.e., photosynthesis per unit leaf area) as the sampling attribute. Ratio and regression estimators with leaf area as an auxiliary variable were also studied. On the basis of their bias and accuracy in estimating instantaneous whole-tree photosynthesis (mg CO(2)), the physiologically based sampling schemes were superior for all combinations of clone type, weather condition and leaf orientation. Aspects of extending the sampling process to estimate daily and seasonal photosynthesis are also elaborated.     

Photosynthetic response of poplar leaves at different developmental phases to environmental factors

With economic incentives and interests in fast-growing poplar trees for short-rotation production of fiber and veneer, many new poplar hybrids have been bred and planted in China, but how to match the new poplar clones to suitable sites and maintain their higher growth rates is still not very clear. In this study, the photosynthetic response of poplar leaves at various developmental stages during two seasons (summer and autumn) was explored and mechanistic models for the photosynthesis of poplar leaves at different developmental phases in response photosynthetic active radiation (PAR), temperature, and relatively humidity were established using the optimization software package 1st Opt. Mature poplar leaves in autumn had significantly higher photosynthetic capacity than leaves at other stages and seasons. Based on the models established for poplar leaves at different phases, the main limiting factors for photosynthesis at the research site were high PAR and temperature in the summer and low PAR in the autumn. Our results highlight the importance of selecting suitable sites, pruning and stand density control during the plantation development to maintain higher photosynthetic rates of poplar trees and to establish optimum cultivation patterns for various utilization of poplar plantations.     

四种杨树扦插苗的硝酸还原酶活力体内衰减的初步研究

硝酸还原酶(NADH:NR E.C.1.6.6.1)是植物体内同化硝酸盐的关键酶,可作为作物育种和营养诊断的生化指标。在木本植物方面,硝酸还原酶的研究正在不断向广度和深度发展。     

田间条件下毛白杨无性系光合和蒸腾特性研究

测定了８个毛白杨无性系净光合速率、蒸腾速率、蒸腾效率和气孔阻力等生理指标，反映出它们抗旱性上的差异。毛白杨无性系净光合速率和蒸腾速率与其叶片形态特征和解剖特征有着密切关系。毛白杨各无性系单位面积日同化量之间的差异，来源于它们在１２：００ｐｍ─２：００ｐｍ时，对逆境忍受能力的差异和时至４：００ｐｍ时叶细胞膨压恢复能力的差异。短时间内测定的净光合速率不能作为预测树木生长量大小的直接指标。     

Leaf senescence and late-season net photosynthesis of sun and shade leaves of overstory sweetgum (Liquidambar styraciflua) grown in elevated and ambient carbon dioxide concentrations

We examined the effects of elevated CO2 concentration ([CO2]) on leaf demography, late-season photosynthesis and leaf N resorption of overstory sweetgum (Liquidambar styraciflua L.) trees in the Duke Forest Free Air CO2 Enrichment (FACE) experiment. Sun and shade leaves were subdivided into early leaves (formed in the overwintering bud) and late leaves (formed during the growing season). Overall, we found that leaf-level net photosynthetic rates were enhanced by atmospheric CO2 enrichment throughout the season until early November; however, sun leaves showed a greater response to atmospheric CO2 enrichment than shade leaves. Elevated [CO2] did not affect leaf longevity, emergence date or abscission date of sun leaves or shade leaves. Leaf number and leaf area per shoot were unaffected by CO2 treatment. A simple shoot photosynthesis model indicated that elevated [CO2] stimulated photosynthesis by 60% in sun shoots, but by only 3% in shade shoots. Whole-shoot photosynthetic rate was more than 12 times greater in sun shoots than in shade shoots. In senescent leaves, elevated [CO2] did not affect residual leaf nitrogen, and nitrogen resorption was largely unaffected by atmospheric CO2 enrichment, except for a small decrease in shade leaves. Overall, elevated [CO2] had little effect on the number of leaves per shoot at any time during the season and, therefore, did not change seasonal carbon gain by extending or shortening the growing season. Stimulation of carbon gain by atmospheric CO2 enrichment in sweetgum trees growing in the Duke Forest FACE experiment was the result of a strong stimulation of photosynthesis throughout the growing season.     

西藏两种杨树光合特性及其释氧价值评估

采用Li-Cor-6400光合仪对藏川杨和北京杨的日光合特性进行测定,结果表明：藏川杨和北京杨的日光合进程均表现为双峰曲线,藏川杨与北京杨的净光合速率日变化最大峰值均在14：00出现,分别为16.62μmol/（m^2·s）和15.52μmol/（m^2·s）;两种杨树的蒸腾速率日变化均呈单峰曲线,峰值均出现在12：00,藏川杨峰值为8.47 mmol/（m^2·s）,北京杨峰值是8.74 mmol/（m^2·s）;单株藏川杨的年释氧生态价值高于北京杨,分别为348元和210元。     

Photosynthetic light response of flooded cherrybark oak (Quercus pagoda) seedlings grown in two light regimes

Two-year-old cherrybark oak (Quercus pagoda Raf.) seedlings raised in full or partial (27%) sunlight were flooded for 30 days to study the effects of light availability and root inundation on photosynthetic light response. Compared with seedlings receiving full sunlight, seedlings receiving partial sunlight developed leaves with 90% greater blade area, 26% less mass per unit volume, and 35% lower nitrogen (N) concentration per unit area, leading to a 15% reduction in leaf photosynthetic capacity when carbon exchange rates were based on blade area. However, when carbon exchange rates were based on leaf mass, leaves acclimated to partial sunlight exhibited a 15% greater photosynthetic capacity realized primarily through an increased initial slope of the photosynthetic light response (A/PPFD) curve and increased net photosynthesis at leaf saturation (Amax). Short-term flooding increased leaf mass per unit area more than 19%, reduced foliar N concentrations per unit dry mass by 19%, and initiated reductions in Amax and apparent quantum yield (phi) of seedlings in both light regimes. Greatest impairment of Amax (56% area basis, 65% mass basis) and phi (40%) were observed in leaves receiving full sunlight, and the declines were concomitant with a 35% decrease in chlorophyll concentration. Flooding also depressed instantaneous photosynthetic N-use efficiency (PPNUE) such that Amax decreased 54%, and the initial slope of PPNUE/PPFD curves decreased 33 and 50% for leaves acclimated to partial and full sunlight, respectively. The A/PPFD patterns indicated that the magnitude of flood-induced inhibition of the photosynthetic mechanism of cherrybark oak seedlings is determined partly by the light environment.     

六个杨树无性系苗木生长、生物量和光合作用的研究*

六个杨树无性系苗木（２年生根１年生于）叶面积、苗高和地径的季生长用Ｒｉｃｈａｒｄｓ方程模拟，根据模拟参数分别计算最大绝对生长速率，平均绝对生长速率，生长期和拐点。结果为：六个无性系之间在生长速率、拐点和生长期方面均有明显的差异；最好的欧美杨６４号生长季末的总生物量是赤峰杨３４号的六倍多；六个无性系的总生物量与它们的总叶面积、平均单个叶片的面积、叶面积的平均生长速率和每株树的日净光合总量等呈正相关，而与单位叶面积的日净光合总量呈负相关；总生物量生产依赖于叶面积、苗高、直径的生长速率及其生长期。     

辽宁省小叶杨叶片解剖学特征研究

小叶杨是我国三北地区重要的耐旱耐瘠薄的乡土杨树。对辽宁省境内的小叶杨进行叶片解剖学研究,结果表明：小叶杨叶片远轴面气孔密度明显高于近轴面,个体间变异较大,与产地没有明显相关性;气孔长度与气孔密度呈极显著负相关;叶片着生部位对小叶杨近轴面气孔密度有明显影响;栅栏组织与海绵组织之比值与耐旱性没有明显相关性。文章同时讨论了国内杨属树种叶片解剖学特点及影响解剖学观测的因素。     

复合经营猕猴桃光合日变化的初步研究

猕猴桃叶片光合速率日变化呈双峰曲线，不同龄叶片具有不同的光合速率。套种植物狗尾草、美洲黑杨和印尼大豆的光合速率日变化也基本呈一双峰曲线。套种可以明显降低中午猕猴桃光合“午睡”，增加光合速率与各环境因素的相关性，从空间上和时间上更充分地利用光能，从而改善猕猴桃冠幕微气候，提高光能利用率。     

Interrelationships among light,photosynthesis and nitrogen in the crown of mature Pinus contorta ssp. latifolia

Scaling leaf-level measurements to estimate carbon gain of entire leaf crowns or canopies requires an understanding of the distribution of photosynthetic capacity and corresponding light microenvironments within a crown. We have compared changes in the photosynthetic light response and nitrogen (N) content (per unit leaf area) of Pinus contorta Dougl. ssp. latifolia Engelm. (lodgepole pine) leaves in relation to their age and light microenvironment. The vertical gradient in integrated daily photosynthetic photon flux density (PPFD) from the upper to the lower crown of lodgepole pine was similar in magnitude to the horizontal gradient in daily PPFD along shoots from young to old leaves. The relationship between light-saturated net photosynthesis (A(max)) and daily PPFD was significant for both young and old leaves. However, old leaves had a lower A(max) than young leaves in a similar daily irradiance regime. For leaves of all ages from throughout the crown, A(max) was linearly related to the estimated daily net carbon gain that leaves could achieve in their natural PPFD environment (estimated A(day)) (r(2) = 0.84, P < 0.001, n = 39), indicating that estimated A(day) may be dominated by carbon fixed when leaves are light-saturated and operating at A(max). Comparison of the PPFD required to achieve A(max) and the PPFD available to the leaves showed that all of the measured leaves (n = 39), regardless of their position in the crown or age, were in light environments that could light-saturate photosynthesis for a similar proportion of the day. For all data pooled, foliar N was weakly correlated with daily PPFD. Analyzing each leaf age class separately showed that foliar N was significantly related to daily PPFD, A(max), and estimated A(day) for the youngest leaves but not for middle-aged or old leaves. Therefore, the general theory that foliar N is allocated within a crown according to total daily light availability was supported only for young (1-4 years old) leaves in this study.     

Resistance to Adversity of New Poplar Clones

1IntroductionAdversitystudyisoneofthefocusesinplantphysiology.However,thepriorityhadbeenputontheselectionofpoplarcloneswithfastgrowthrateandbetterwoodqualityonpoplarbreedinginthepastfewdecades.Therewerelimitedresearchestargetedontheresistancebreeding.Moreover,mostoftheresearchonadversityresistancewasconductedonPopulustomentosa(Peietal.1994,Duanetal.1998)andPopulusussuriensis(Jiangetal.1999).LittlewasknownabouttheadversityresistanceofpoplarclonesinSectionAigeiros(Wang1982,Zhu1990,Liuetal.1…     

Diversity of leaf traits related to productivity in 31 Populus deltoides x Populus nigra clones

To test if some leaf parameters are predictors of productivity in a range of Populus deltoides (Bartr.) Marsh. x P. nigra L. clones, we assessed leaf traits and productivity in 2-month-old rooted cuttings from 31 clones growing in 4-l pots in a greenhouse, under conditions of controlled temperature and optimal irrigation. We evaluated four groups of variables describing (1) productivity (total biomass), (2) leaf growth (total leaf number increment and total leaf area increment rate), (3) leaf structure (specific leaf area and nitrogen and carbon contents) and (4) carbon isotope discrimination (delta), which is negatively correlated with time-integrated water-use efficiency. High-yielding clones did not necessarily display high leaf growth rates, but they displayed a larger total leaf area, lower specific leaf area and lower leaf nitrogen concentration than clones with low productivity. Total leaf area was mainly controlled by maximal individual leaf area and total leaf area increment rate (r = 0.51 and 0.56, respectively). Carbon isotope discrimination did not correlate with total biomass, but it was associated with total number of leaves and total leaf area increment rate (r = 0.39 and 0.45, respectively). Therefore, leaf area and specific leaf area were better indicators of productivity than leaf growth traits. The observed independence of delta from biomass production provides opportunities for selecting poplar clones combining high productivity and high water-use efficiency.