GENETIC VARIANCE AND COVARIANCE FOR PHYSIOLOGICAL TRAITS IN LOBELIA:ARE THERE CONSTRAINTS ON ADAPTIVE EVOLUTION?

Physiological traits that control the uptake of carbon dioxide and loss of water are key determinants of plant growth and reproduction. Variation in these traits is often correlated with environmental gradients of water, light, and nutrients, suggesting that natural selection is the primary evolutionary mechanism responsible for physiological diversification. Responses to selection, however, can be constrained by the amount of standing genetic variation for physiological traits and genetic correlations between these traits. To examine the potential for constraint on adaptive evolution, we estimated the quantitative genetic basis of physiological trait variation in one population of each of two closely related species (Lobelia siphilitica and L. cardinalis). Restricted maximum likelihood analyses of greenhouse-grown half-sib families were used to estimate genetic variances and covariances for seven traits associated with carbon and water relations. We detected significant genetic variation for all traits in L. siphilitica, suggesting that carbon-gain and water-use traits could evolve in response to natural selection in this population. In particular, narrow-sense heritabilities for photosynthetic rate (A), stomatal conductance (gs), and water-use efficiency (WUE) in our L. siphilitica population were high relative to previous studies in other species. Although there was significant narrow-sense heritability for A in L. cardinalis, we detected little genetic variation for traits associated with water use (gs and WUE), suggesting that our population of this species may be unable to adapt to drier environments. Despite being tightly linked functionally, the genetic correlation between A and gs was not strong and significant in either population. Therefore, our L. siphilitica population would not be genetically constrained from evolving high A (and thus fixing more carbon for growth and reproduction) while also decreasing gs to limit water loss. However, a significant negative genetic correlation existed between WUE and plant size in L. siphilitica, suggesting that high WUE may be negatively associated with high fecundity. In contrast, our results suggest that any constraints on the evolution of photosynthetic and stomatal traits of L. cardinalis are caused primarily by a lack of genetic variation, rather than by genetic correlations between these functionally related traits.     

Photosynthetic pathway and ecological adaptation explain stomatal trait diversity amongst grasses

? The evolution of C(4) photosynthesis in plants has allowed the maintenance of high CO(2) assimilation rates despite lower stomatal conductances. This underpins the greater water-use efficiency in C(4) species and their tendency to occupy drier, more seasonal environments than their C(3) relatives. ? The basis of interspecific variation in maximum stomatal conductance to water (g(max) ), as defined by stomatal density and size, was investigated in a common-environment screening experiment. Stomatal traits were measured in 28 species from seven grass lineages, and comparative methods were used to test for predicted effects of C(3) and C(4) photosynthesis, annual precipitation and habitat wetness on g(max) . ? Novel results were as follows: significant phylogenetic patterns exist in g(max) and its determinants, stomatal size and stomatal density; C(4) species consistently have lower g(max) than their C(3) relatives, associated with a shift towards smaller stomata at a given density. A direct relationship between g(max) and precipitation was not supported. However, we confirmed associations between C(4) photosynthesis and lower precipitation, and showed steeper stomatal size-density relationships and higher g(max) in wetter habitats. ? The observed relationships between stomatal patterning, photosynthetic pathway and habitat provide a clear example of the interplay between anatomical traits, physiological innovation and ecological adaptation in plants.     

Trait convergence and plasticity among native and invasive species in resource-poor environments

? Premise of study: Functional trait comparisons provide a framework with which to assess invasion and invasion resistance. However, recent studies have found evidence for both trait convergence and divergence among coexisting dominant native and invasive species. Few studies have assessed how multiple stresses constrain trait values and plasticity, and no study has included direct measurements of nutrient conservation traits, which are critical to plants growing in low-resource environments. ? Methods: We evaluated how nutrient and water stresses affect growth and allocation, water potential and gas exchange, and nitrogen (N) allocation and use traits among a suite of six codominant species from the Intermountain West to determine trait values and plasticity. In the greenhouse, we grew our species under a full factorial combination of high and low N and water availability. We measured relative growth rate (RGR) and its components, total biomass, biomass allocation, midday water potential, photosynthetic rate, water-use efficiency (WUE), green leaf N, senesced leaf N, total N pools, N productivity, and photosynthetic N use efficiency. ? Key results: Overall, soil water availability constrained plant responses to N availability and was the major driver of plant trait variation in our analysis. Drought decreased plant biomass and RGR, limited N conservation, and led to increased WUE. For most traits, native and nonnative species were similarly plastic. ? Conclusions: Our data suggest native and invasive biomass dominants may converge on functionally similar traits and demonstrate comparable ability to respond to changes in resource availability.     

Leaf Gas Exchange of Pachyrhizus ahipa and P. erosus Under Water and Temperature Stress

Gas exchange, water relations, and leaf traits were studied in the tuberous-root producing legumes ahipa (Pachyrhizus ahipa) and yambean (P. erosus) under different environmental conditions. Differences in leaf traits (hairiness, leaf area, areal leaf mass, stomatal density) and paraheliotropism were found between ahipa and yambean. Under sufficient water supply, the increase in air temperature and decrease in air humidity increased stomatal conductance (g _s) and net photosynthetic rate (P _N) in yambean but reduced them in ahipa. In a drying soil (14 d after irrigation), inter-specific variation in gas exchange was only observed in the early morning, and yambean showed a greater sensitivity to water restriction than ahipa. High g _s at low humidity increased P _N of P. erosus but resulted in lower water-use efficiency (WUE). However, long-term WUE, estimated by leaf carbon isotope discrimination, showed little variation between species. Daily-irrigated ahipa and yambean grown in the greenhouse did not show significant differences in gas exchange. However, leaf temperature was significantly greater in yambean than in ahipa while a steepper relationship between E and P _N and g _s was observed in ahipa.     

Photosynthetic response of four fern species from different habitats to drought stress: relationship between morpho-anatomical and physiological traits

Ferns flourish in many habitats, from epiphytic to terrestrial and from sunny to shady, and such varied conditions require contrasting photosynthetic strategies to cope with drought. Four species of temperate ferns from different habitats were subjected to drought by withholding irrigation in order to investigate their photosynthetic responses. Lepisorus thunbergianus (epiphytic) had low stomatal density and showed high water-use efficiency (WUE) retaining photosynthetic activity with low relative frond water content under drought stress, which suggested their high adaptation to drought. On the other hand, low WUE with low light-saturated photosynthetic rate in Adiantum pedatum (terrestrial, shady environment) was associated with much lower photosynthesis than in the other species under drought stress, suggesting lower adaptation to drought-prone habitats. Morphological stomatal traits such as stomatal density and photosynthetic response to drought in ferns involved species-specific adaptation to survive and grow in their natural habitats with different levels of drought.     

Quantitative variation in water-use efficiency across water regimes and its relationship with circadian, vegetative, reproductive, and leaf gas-exchange traits

Drought limits light harvesting, resulting in lower plant growth and reproduction. One trait important for plant drought response is water-use efficiency (WUE). We investigated (1) how the joint genetic architecture of WUE, reproductive characters, and vegetative traits changed across drought and well-watered conditions, (2) whether traits with distinct developmental bases (e.g. leaf gas exchange versus reproduction) differed in the environmental sensitivity of their genetic architecture, and (3) whether quantitative variation in circadian period was related to drought response in Brassica rapa. Overall, WUE increased in drought, primarily because stomatal conductance, and thus water loss, declined more than carbon fixation. Genotypes with the highest WUE in drought expressed the lowest WUE in well-watered conditions, and had the largest vegetative and floral organs in both treatments. Thus, large changes in WUE enabled some genotypes to approach vegetative and reproductive trait optima across environments. The genetic architecture differed for gas-exchange and vegetative traits across drought and well-watered conditions, but not for floral traits. Correlations between circadian and leaf gas-exchange traits were significant but did not vary across treatments, indicating that circadian period affects physiological function regardless of water availability. These results suggest that WUE is important for drought tolerance in Brassica rapa and that artificial selection for increased WUE in drought will not result in maladaptive expression of other traits that are correlated with WUE.     

两种生态型榕树的叶绿素含量、荧光特性和叶片气体交换日变化的比较研究

比较盆栽 生榕树和两栖型树的形态差异、叶片叶绿素含量、叶绿素荧光特性和气体交换的日变化。两栖型榕树具有较发达的气生根和水生不定根,叶片比陆生榕树宽,并有向中生性 倾向,陆生榕树的叶绿素含量比两栖榕树高,净光合速率略高于水培两栖型榕树,但明显高于土培两栖型榕树,蒸腾速率以水培两栖型树最高,陆生榕树次之,土培两栖型榕树最低,线性回归分析表明,三者的叶片气孔导度与净光合速率变化均呈正相关,气孔导度的变化     

Genetic variation in and covariation between leaf gas exchange,morphology, and development in Polygonum arenastrum,an annual plant

Summary We present evidence of genetic variation in and covariation between leaf-level gas exchange properties and leaf size among family lines of Polygonum arenastrum. This self-fertilizing annual had previously been shown to vary genetically in developmental phenology and in morphology (size of leaves, internodes, flowers and seeds) (Geber 1990). Significant family differences were found in photosynthetic carbon assimilation rate (A), lcaf conductance to water vapor (g), instantaneous water-use efficiency (WUE), and leaf carbon isotope discrimination (). A strong positive genetic correlation between A and g suggested that there was stomatal limitation on A. In addition, higher g led to relatively greater increases in transpiration, E, than in assimilation, A, so that families with high rates of gas exchange had lower instantaneous WUE and/or higher carbon isotope discrimination values. Leaf size and gas exchange were genetically correlated. In earlier studies leaf size was found to be genetically correlated with developmental phenology (Geber 1990). The pattern that emerges is one in which small-leaved families (which also have small internodes, flowers, and seeds) tend to have high gas exchange rates, low WUE, rapid development to flowering and high early fecundity, but reduced life span and maximum (vegetative and reproductive) yield compared to large-leaved families. We suggest that this pattern may have arisen from selection for contrasting suites of characters adapted to environments differing in season length.     

Stomatal kinetics and photosynthetic gas exchange along a continuum of isohydric to anisohydric regulation of plant water status

Species' differences in the stringency of stomatal control of plant water potential represent a continuum of isohydric to anisohydric behaviours. However, little is known about how quasi‐steady‐state stomatal regulation of water potential may relate to dynamic behaviour of stomata and photosynthetic gas exchange in species operating at different positions along this continuum. Here, we evaluated kinetics of light‐induced stomatal opening, activation of photosynthesis and features of quasi‐steady‐state photosynthetic gas exchange in 10 woody species selected to represent different degrees of anisohydry. Based on a previously developed proxy for the degree of anisohydry, species' leaf water potentials at turgor loss, we found consistent trends in photosynthetic gas exchange traits across a spectrum of isohydry to anisohydry. More anisohydric species had faster kinetics of stomatal opening and activation of photosynthesis, and these kinetics were closely coordinated within species. Quasi‐steady‐state stomatal conductance and measures of photosynthetic capacity and performance were also greater in more anisohydric species. Intrinsic water‐use efficiency estimated from leaf gas exchange and stable carbon isotope ratios was lowest in the most anisohydric species. In comparisons between gas exchange traits, species rankings were highly consistent, leading to species‐independent scaling relationships over the range of isohydry to anisohydry observed.     

Control of water-use efficiency by florigen

A major issue in modern agriculture is water loss through stomata during photosynthetic carbon assimilation. In water-limited ecosystems, annual plants have strategies to synchronize their growth and reproduction to the availability of water. Some species or ecotypes of flowers are early to ensure that their life cycles are completed before the onset of late season terminal drought (“drought escape”). This accelerated flowering correlates with low water-use efficiency (WUE). The molecular players and physiological mechanisms involved in this coordination are not fully understood. We analyzed WUE using gravimetry, gas exchange, and carbon isotope discrimination in florigen deficient (sft mutant), wild-type (Micro-Tom), and florigen over-expressing (SFT-ox) tomato lines. Increased florigen expression led to accelerated flowering time and reduced WUE. The low WUE of SFT-ox was driven by higher stomatal conductance and thinner leaf blades. This florigen-driven effect on WUE appears be independent of abscisic acid (ABA). Our results open a new avenue to increase WUE in crops in an ABA-independent manner. Manipulation of florigen levels could allow us to produce crops with a life cycle synchronized to water availability.     

Fine root dynamics in relation to nutrients in oligotrophic Bornean rain forest soils

The fine roots of plants are key structures enabling soil resource acquisition, yet our understanding of their dynamics and the factors governing them is still underdeveloped, especially in tropical forests. We evaluated whether Bornean tree communities on soils with contrasting resource availability display different soil resource uptake strategies, based on their fine root properties and dynamics, and related responses of fine roots to the availability of multiple nutrients. Using root cores and ingrowth cores, we quantified variation in community-level fine root properties (biomass, length, and area) and their growth rates, biomass turnover rate, and specific root length (SRL) between clay and sandy loam soils, on which tree community composition differs dramatically. We found that standing fine root biomass and biomass, length, and area growth were higher in sandy loam, the soil type that is better-drained, coarser-textured, and less fertile for most nutrients. In clay SRL was significantly greater, and turnover tended to be faster, than in sandy loam. Across both soils, greater supplies of K⁺, NH₄ ⁺, and PO₄ ^3? were associated with greater standing biomass and growth rates of fine roots, suggesting foraging for these nutrients. Our data support the hypothesis that the sandy loam tree community achieves fine root absorptive area through faster growth and greater investment on a mass basis, whereas trees on clay achieve a similar standing absorptive area through slower growth of less-dense fine root tissues. Furthermore, our results suggest colimitation by multiple nutrients, which may enhance tree species coexistence through increased dimensionality of soil-resource niches.     

Mechanisms of selection for drought stress tolerance and avoidance in Impatiens capensis (Balsaminaceae)

For longer lived annual plants, high water-use efficiency (WUE) and low stomatal conductance are hypothesized to confer a fitness advantage under drought stress. To directly test the adaptive significance of WUE and stomatal conductance under drought stress, inbred lines of Impatiens capensis were grown in two field environments (watered and not-watered), in a year of unusual early-season drought. In contrast to the results from a previous study of late-season drought in the same system, selection was detected for lower WUE, increased stomatal conductance, and early flowering time. These findings suggest that early-season drought conditions may select for drought avoidance traits such as low WUE and early reproduction, whereas later drought selects for tolerance traits such as high WUE.     

Correlated evolution of stem and leaf hydraulic traits in Pereskia (Cactaceae)

Recent studies have demonstrated significant correlations between stem and leaf hydraulic properties when comparing across species within ecological communities. This implies that these traits are co-evolving, but there have been few studies addressing plant water relations within an explicitly evolutionary framework. This study tests for correlated evolution among a suite of plant water-use traits and environmental parameters in seven species of Pereskia (Cactaceae), using phylogenetically independent contrasts. There were significant evolutionary correlations between leaf-specific xylem hydraulic conductivity, Huber Value, leaf stomatal pore index, leaf venation density and leaf size, but none of these traits appeared to be correlated with environmental water availability; only two water relations traits - mid-day leaf water potentials and photosynthetic water use efficiency - correlated with estimates of moisture regime. In Pereskia, it appears that many stem and leaf hydraulic properties thought to be critical to whole-plant water use have not evolved in response to habitat shifts in water availability. This may be because of the extremely conservative stomatal behavior and particular rooting strategy demonstrated by all Pereskia species investigated. These results highlight the need for a lineage-based approach to understand the relative roles of functional traits in ecological adaptation.     

Light and VPD gradients drive foliar nitrogen partitioning and photosynthesis in the canopy of European beech and silver fir

While foliar photosynthetic relationships with light, nitrogen, and water availability have been well described, environmental factors driving vertical gradients of foliar traits within forest canopies are still not well understood. We, therefore, examined how light availability and vapour pressure deficit (VPD) co-determine vertical gradients (between 12 and 42 m and in the understorey) of foliar photosynthetic capacity (Amax), 13C fractionation (∆), specific leaf area (SLA), chlorophyll (Chl), and nitrogen (N) concentrations in canopies of Fagus sylvatica and Abies alba growing in a mixed forest in Switzerland in spring and summer 2017. Both species showed lower Chl/N and lower SLA with higher light availability and VPD at the top canopy. Despite these biochemical and morphological acclimations, Amax during summer remained relatively constant and the photosynthetic N-use efficiency (PNUE) decreased with higher light availability for both species, suggesting suboptimal N allocation within the canopy. ∆ of both species were lower at the canopy top compared to the bottom, indicating high water-use efficiency (WUE). VPD gradients strongly co-determined the vertical distribution of Chl, N, and PNUE in F. sylvatica, suggesting stomatal limitation of photosynthesis in the top canopy, whereas these traits were only related to light availability in A. alba. Lower PNUE in F. sylvatica with higher WUE clearly indicated a trade-off in water vs. N use, limiting foliar acclimation to high light and VPD at the top canopy. Species-specific trade-offs in foliar acclimation to environmental canopy gradients may thus be considered for scaling photosynthesis from leaf to canopy to landscape levels.     

Carbon isotope discrimination and stomatal responses of mature Pinus sylvestris L. trees exposed in situ for three years to elevated CO2 and temperature

The Climate Change Experiment (CLIMEX) is a unique large scale facility in which an entire undisturbed catchment of boreal vegetation has been exposed to elevated CO₂ (560 ppm) and temperature (+3°C summer, +5°C winter) for the past three years with all the soil-plant-atmosphere linkages intact. Here, carbon isotope composition and stomatal density have been analysed from sequential year classes of needles of mature Scots pine trees (Pinus sylvestris L.) to investigate the response of time-integrated water-use efficiency (UWE) and stomatal density to CO₂ enrichment and climate change. Carbon isotope discrimination decreased and WUE increased in cohorts of needles developing under increased CO₂ and temperature, compared to needles on the same trees developing in pretreatment years. Mid-season instantaneous gas exchange, measured on the same trees for the past four years, indicated that these responses resulted from higher needle photosynthetic rates and reduced stomatal conductance. Needles of P. sylvestris developing under increased CO₂ and temperature had consistently lower stomatal densities than their ambient grown counterparts on the same trees. The stomatal density of P. sylvestris needles was inversely correlated with δ¹³C-derived WUE, implying some effect of this morphological response on leaf gas exchange. Future atmospheric CO₂ and temperature increases are therefore likely to improve the water economy of P. sylvestris, at least at the scale of individual needles, by affecting stomatal density and gas exchange processes.     

Specific leaf area and leaf nitrogen concentration in annual and perennial grass species growing in Mediterranean old-fields

 We evaluated the hypothesis that photosynthetic traits differ between leaves produced at the beginning (May) and the end (November–December) of the rainy season in the canopy of a seasonally dry forest in Panama. Leaves produced at the end of the wet season were predicted to have higher photosynthetic capacities and higher water-use efficiencies than leaves produced during the early rainy season. Such seasonal phenotypic differentiation may be adaptive, since leaves produced immediately preceding the dry season are likely to experience greater light availability during their lifetime due to reduced cloud cover during the dry season. We used a construction crane for access to the upper canopy and sampled 1- to 2-month-old leaves marked in monthly censuses for six common tree species with various ecological habits and leaf phenologies. Photosynthetic capacity was quantified as light- and CO₂-saturated oxygen evolution rates with a leaf-disk oxygen electrode in the laboratory (O_2max) and as light-saturated CO₂ assimilation rates of intact leaves under ambient CO₂ (A_max). In four species, pre-dry season leaves had significantly higher leaf mass per unit area. In these four species, O_2max and A_max per unit area and maximum stomatal conductances were significantly greater in pre-dry season leaves than in early wet season leaves. In two species, A_max for a given stomatal conductance was greater in pre-dry season leaves than in early wet season leaves, suggesting a higher photosynthetic water-use efficiency in the former. Photosynthetic capacity per unit mass was not significantly different between seasons of leaf production in any species. In both early wet season and pre-dry season leaves, mean photosynthetic capacity per unit mass was positively correlated with nitrogen content per unit mass both within and among species. Seasonal phenotypic differentiation observed in canopy tree species is achieved through changes in leaf mass per unit area and increased maximum stomatal conductance rather than by changes in nitrogen allocation patterns. Received: 7 March 1996 / Accepted: 1 August 1996     

Independent variation in photosynthetic capacity and stomatal conductance leads to differences in intrinsic water use efficiency in 11 soybean genotypes before and during mild drought

Intrinsic water use efficiency (WUE(intr)), the ratio of photosynthesis to stomatal conductance to water, is often used as an index for crop water use in breeding projects. However, WUE(intr) conflates variation in these two processes, and thus may be less useful as a selection trait than knowledge of both components. The goal of the present study was to determine whether the contribution of photosynthetic capacity and stomatal conductance to WUE(intr) varied independently between soybean genotypes and whether this pattern was interactive with mild drought. Photosynthetic capacity was defined as the variation in WUE(intr) that would occur if genotypes of interest had the same stomatal conductance as a reference genotype and only differed in photosynthesis; similarly, the contribution of stomatal conductance to WUE(intr) was calculated assuming a constant photosynthetic capacity across genotypes. Genotypic differences in stomatal conductance had the greatest effect on WUE(intr) (26% variation when well watered), and was uncorrelated with the effect of photosynthetic capacity on WUE(intr). Thus, photosynthetic advantages of 8.3% were maintained under drought. The maximal rate of Rubisco carboxylation, generally the limiting photosynthetic process for soybeans, was correlated with photosynthetic capacity. As this trait was not interactive with leaf temperature, and photosynthetic capacity differences were maintained under mild drought, the observed patterns of photosynthetic advantage for particular genotypes are likely to be consistent across a range of environmental conditions. This suggests that it is possible to employ a selection strategy of breeding water-saving soybeans with high photosynthetic capacities to compensate for otherwise reduced photosynthesis in genotypes with lower stomatal conductance.     

Genetic variation of stomatal traits and carbon isotope discrimination in two hybrid poplar families (Populus deltoides 'S9-2' x P. nigra 'Ghoy' and P. deltoides 'S9-2' x P. trichocarpa 'V24')

Background and Aims

Stomata play an important role in both the CO₂ assimilation and water relations of trees. Therefore, stomatal traits have been suggested as criteria for selection of clones or genotypes which are more productive and have larger water-use efficiency (WUE) than others. However, the relationships between plant growth, WUE and stomatal traits are still unclear depending on plant material (genus, species, families, genotypes) and, more precisely, on the strength of the relationships between the plants. In this study, the correlations between these three traits categories, i.e. plant growth, WUE and stomatal traits, were compared in two related poplar families.

Methods

Stomatal traits (stomatal density, length and ratio adaxial : abaxial stomatal densities) of a selection of F₁ genotypes and the parents of two hybrid poplar families Populus deltoides ‘S9-2’ × P. nigra ‘Ghoy’ (D × N family, 50 F₁) and P. deltoides ‘S9-2’ × P. trichocarpa ‘V24’ (D × T family, 50 F₁) were measured, together with stem height and circumference. Carbon isotope discrimination (Δ) was determined and used as an indicator of leaf-level intrinsic WUE.

Key Results

Leaves of hybrids and parents were amphistomatous, except for the P. trichocarpa parent. Both families displayed high values of heritability for stomatal traits and Δ. In the progeny, the relationship between stem circumference and Δ was weak for the D × N family, while abaxial and total stomatal density were positively associated with stem dimensions for the D × T family only.

Conclusions

Genetic variation in stomatal traits and Δ was large within as well as between the different poplar species and their hybrids, but there were no direct relationships between stomatal traits and plant growth or Δ. As already noticed in various poplar hybrids, the absence of, or the weak, relationship between Δ and plant growth allows the possibility of selecting poplar genotypes combining high productivity and high WUE. In this study, stomatal traits are of limited value as criteria for selection of genotypes with good growth and large WUE.Key words: Adaxial and abaxial stomatal density, stomatal length, heritability, water-use efficiency (WUE), F₁ hybrids, breeding, Populus deltoides, Populus nigra, Populus trichocarpa     

彰武松、樟子松光合生产与蒸腾耗水特性

本文采用Li-6400光合测定系统对性成熟(18a)阶段彰武松(Pinus densiflora var.zhangwuensis)和樟子松(Pinus sylvestris var.mongolica)光合及蒸腾指标不同季节日变化进行了测定,并采用切枝蒸腾法对两个树种叶片气孔蒸腾和角质层蒸腾进行对比测定,评价了气孔开闭敏感性,探讨了两个树种光合生产与蒸腾耗水特性。结果表明：在同样生境条件下,彰武松比樟子松有较大的光合速率(Pn)和较小的蒸腾速率(Tr)。在5月和7月,彰武松的Pn和Tr日变化呈现明显双峰型,其Pn和Tr“午休”现象均主要受气孔限制;在10月呈单峰型。樟子松的Pn和Tr日变化在整个生长季均呈单峰型,而且,彰武松日光合量(DAP)均高于樟子松,是樟子松的163.4%(5月)、211.1%(7月)和183.6%(10月)。光响应曲线参数表明：在不同月份,彰武松最大光合速率(Pmax)均大于樟子松,且光饱和点(LSP)较高,光补偿点(LCP)较低。在任意被测时刻,彰武松气孔导度(Gs)和Tr都小于樟子松。彰武松具有较小气孔和角质层蒸腾速度,并且在同样干旱条件下,彰武松气孔下陷,其气孔的开闭反应更加敏感。彰武松水分利用效率(WUE)较高,约是樟子松的2.29倍。这些结果暗示,彰武松以其高的光合速率和低的蒸腾耗水特性,提高水分利用效率,以其敏感的气孔开闭机制和旱生叶片结构进而实现在干旱半干旱地区的速生特性。