Environmental and stomatal control of photosynthetic enhancement in the canopy of a sweetgum (Liquidambar styraciflua L.) plantation during 3 years of CO2 enrichment

Light‐saturated photosynthetic and stomatal responses to elevated CO₂ were measured in upper and mid‐canopy foliage of a sweetgum (Liquidambar styraciflua L) plantation exposed to free‐air CO₂ enrichment (FACE) for 3 years, to characterize environmental interactions with the sustained CO₂ effects in an intact deciduous forest stand. Responses were evaluated in relation to one another, and to seasonal patterns and natural environmental stresses, including high  temperatures, vapour pressure deficits (VPD), and drought. Photosynthetic CO₂ assimilation (A) averaged 46% higher in the +200 µmol mol^?1 CO₂ treatment, in mid‐ and upper canopy foliage. Stomatal conductance (g_s) averaged 14% (mid‐canopy) and 24% (upper canopy) lower under CO₂ enrichment. Variations in the relative responses of A and g_s were linked, such that greater relative stimulation of A was observed on dates when relative reductions in g_s were slight. Dry soils and high VPD reduced g_s and A in both treatments, and tended to diminish treatment differences. The absolute effects of CO₂ on A and g_s were minimized whenever g_s was low (<0·15 mol m^?2 s^?1), but relative effects, as the ratio of elevated to ambient rates, varied greatly under those conditions. Both stomatal and non‐stomatal limitations of A were involved during late season droughts. Leaf temperature had a limited influence on A and g_s, and there was no detectable relationship between prevailing temperature and CO₂ effects on A or g_s. The responsiveness of A and g_s to elevated CO₂, both absolute and relative, was maintained through time and within the canopy of this forest stand, subject to seasonal constraints and variability associated with limiting air and soil moisture.     

鄂东南低山丘陵区枫香种群的天然更新方式与特征

鄂东南低山丘陵区是一个生态脆弱区,枫香（Liquidam bar formasana）林是该地区的重要森林植被类型,对于维持该地区的生态安全和经济发展具有重要意义。为了探明枫香实生和萌芽更新对枫香种群维持的作用,采用样方法研究了该地区枫香林和马尾松（Pinusm assoniana）-枫香混交林枫香种群的天然更新方式,并采用不同年代伐桩构建的时间序列研究了枫香伐桩的萌芽更新规律。结果表明：（1）该地区的枫香种群由实生和萌芽更新植株组成,但二者的比例在马尾松-枫香混交林和枫香林间存在明显差异;（2）萌芽更新可以划分为幼苗萌芽和伐桩萌芽更新2类,其中幼苗萌芽更新是对自然环境的适应,伐桩萌芽更新是对人类砍伐的适应;（3）枫香伐桩萌芽更新存在明显的自疏现象,并最终形成1-2杆的萌代主;与实生幼苗相比,伐桩萌条具有生长迅速的特点。上述结果表明,枫香的实生和萌芽更新共同维持了该地区枫香种群的稳定。     

Leaf dynamics of a deciduous forest canopy: no response to elevated CO<Subscript>2</Subscript>

Leaf area index (LAI) and its seasonal dynamics are key determinants of terrestrial productivity and, therefore, of the response of ecosystems to a rising atmospheric CO₂ concentration. Despite the central importance of LAI, there is very little evidence from which to assess how forest LAI will respond to increasing [CO₂]. We assessed LAI and related leaf indices of a closed-canopy deciduous forest for 4 years in 25-m-diameter plots that were exposed to ambient or elevated CO₂ (542 ppm) in a free-air CO₂ enrichment (FACE) experiment. LAI of this Liquidambar styraciflua (sweetgum) stand was about 6 and was relatively constant year-to-year, including the 2 years prior to the onset of CO₂ treatment. LAI throughout the 1999–2002 growing seasons was assessed through a combination of data on photosynthetically active radiation (PAR) transmittance, mass of litter collected in traps, and leaf mass per unit area (LMA). There was no effect of [CO₂] on any expression of leaf area, including peak LAI, average LAI, or leaf area duration. Canopy mass and LMA, however, were significantly increased by CO₂ enrichment. The hypothesized connection between light compensation point (LCP) and LAI was rejected because LCP was reduced by [CO₂] enrichment only in leaves under full sun, but not in shaded leaves. Data on PAR interception also permitted calculation of absorbed PAR (APAR) and light use efficiency (LUE), which are key parameters connecting satellite assessments of terrestrial productivity with ecosystem models of future productivity. There was no effect of [CO₂] on APAR, and the observed increase in net primary productivity in elevated [CO₂] was ascribed to an increase in LUE, which ranged from 1.4 to 2.4 g MJ^–1. The current evidence seems convincing that LAI of non-expanding forest stands will not be different in a future CO₂-enriched atmosphere and that increases in LUE and productivity in elevated [CO₂] are driven primarily by functional responses rather than by structural changes. Ecosystem or regional models that incorporate feedbacks on resource use through LAI should not assume that LAI will increase with CO₂ enrichment of the atmosphere.     

Photosynthesis and isoprene emission from trees along an urban–rural gradient in Texas

Isoprene emission is an important mechanism for improving the thermotolerance of plant photosystems as temperatures increase. In this study, we measured photosynthesis and isoprene emission in trees along an urban–rural gradient that serves as a proxy for climate change, to understand daily and seasonal responses to changes in temperature and other environmental variables. Leaf‐level gas exchange and basal isoprene emission of post oak (Quercus stellata) and sweet gum (Liquidambar styraciflua) were recorded at regular intervals over an entire growing season at urban, suburban, and rural sites in eastern Texas. In addition, the temperature and atmospheric carbon dioxide concentration experienced by leaves were experimentally manipulated in spring, early summer, and late summer. We found that trees experienced lower stomatal conductance and photosynthesis and higher isoprene emission, at the urban and suburban sites compared to the rural site. Path analysis indicated a daily positive effect of isoprene emission on photosynthesis, but unexpectedly, higher isoprene emission from urban trees was not associated with improved photosynthesis as temperatures increased during the growing season. Furthermore, urban trees experienced relatively higher isoprene emission at high CO₂ concentrations, while isoprene emission was suppressed at the other sites. These results suggest that isoprene emission may be less beneficial in urban, and potentially future, environmental conditions, particularly if higher temperatures override the suppressive effects of high CO₂ on isoprene emission. These are important considerations for modeling future biosphere–atmosphere interactions and for understanding tree physiological responses to climate change.     

Coordination of anthocyanin decline and photosynthetic maturation in juvenile leaves of three deciduous tree species

Juvenile leaves in high-light environments commonly appear red as a result of anthocyanin pigments, which play a photoprotective role during light-sensitive ontogenetic stages. The loss of anthocyanin during leaf development presumably corresponds to a decreased need for photoprotection, as photosynthetic maturation allows leaves to utilize higher light intensities. However, the relationship between photosynthetic development and anthocyanin decline has yet to be quantitatively described. In this study, anthocyanin concentration was measured against photopigment content, lamina thickness, anatomical development, and photosynthetic CO(2) exchange in developing leaves of three deciduous tree species. In all species, anthocyanin disappearance corresponded with development of c. 50% mature photopigment concentrations, c. 80% lamina thickness, and differentiation of the mesophyll into palisade and spongy layers. Photosynthetic gas exchange correlated positively with leaf thickness and chlorophyll content, and negatively with anthocyanin concentration. Species with more rapid photosynthetic maturation lost anthocyanin earliest in development. Chlorophyll a/b ratios increased with leaf age, and were lower than those of acyanic species, consistent with a shading effect of anthocyanin. These results suggest that anthocyanin reassimilation is linked closely with chloroplast and whole-leaf developmental processes, supporting the idea that anthocyanins protect tissues until light processing and carbon fixation have matured to balance energy capture with utilization.     

不同光环境下枫香幼苗的叶片解剖结构

落叶阔叶树种枫香是常绿阔叶林的优势种之一。对比研究了重庆丰都世坪林场不同光环境下枫香幼苗叶片的解剖特征,结果表明:1)旷地枫香叶片的厚度大,气孔个体较大且排列紧密,栅栏组织发达,具有明显的旱生叶特征;而林下的枫香叶片明显变薄,气孔小且排列稀疏,海绵组织发达,具有一定的阴生叶特征;2)不同光环境下,气孔密度、栅栏组织的厚度和层数、栅栏细胞长度以及栅栏组织与海绵组织厚度比(P/S)等具有较高的可塑性,表明这些因素对于枫香适应不同光环境具有较大的作用;3)从叶片解剖特征与功能的统一来看,阳性树种枫香幼苗更适合生长在旷地等高光生境中,而不易生长在郁闭的林下;4)落叶阔叶树种在中亚热带常绿阔叶林群落恢复与演替中具有重要作用。     

枫香叶采后挥发物表征及影响因子研究

以枫香(Liquidambar formosana)叶为材料,采用气相色谱/质谱(GC/MS)联用技术对不同发育程度、干燥时间、储存方式及萃取温度下枫香叶挥发性成分进行分析,以揭示枫香叶释放挥发性有机化合物数量及不同采后处理下挥发物的消长规律。结果表明：(1)枫香叶释放的萜烯类化合物以α-蒎烯、β-蒎烯、β-水芹烯、柠檬烯为主;(2)嫩叶挥发性成分种类较成熟叶高,莰烯、β-萜品烯、罗勒烯仅存在于嫩叶,3-蒈烯、γ-松油烯仅存在于成熟叶;(3)干燥时间对枫香叶挥发性化合物影响较大,挥发物种类随干燥时间延长呈逐渐下降趋势,但主要的萜烯类化合物在不同干燥时间内均能被检测到,表现相对稳定;(4) –5 ℃及17 ℃储存可保留更多挥发性化合物,室温保存则失去较多化合物,但主要的萜烯类化合物在不同储存方式下均可被检测到;(5) 90 ℃高温萃取可获得更多的挥发性化合物,但70 ℃萃取得到的萜烯类化合物相对含量较高,室温萃取所得化合物种类虽最少,但更能反映园林应用中枫香叶芳香物质挥发的实际情况。研究结果可为枫香叶的采后处理及高值化资源利用提供参考。     

Soil carbon and nitrogen cycling and storage throughout the soil profile in a sweetgum plantation after 11 years of CO2‐enrichment

Increased partitioning of carbon (C) to fine roots under elevated [CO₂], especially deep in the soil profile, could alter soil C and nitrogen (N) cycling in forests. After more than 11 years of free‐air CO₂ enrichment in a Liquidambar styraciflua L. (sweetgum) plantation in Oak Ridge, TN, USA, greater inputs of fine roots resulted in the incorporation of new C (i.e., C with a depleted δ¹³C) into root‐derived particulate organic matter (POM) pools to 90‐cm depth. Even though production in the sweetgum stand was limited by soil N availability, soil C and N contents were greater throughout the soil profile under elevated [CO₂] at the conclusion of the experiment. Greater C inputs from fine‐root detritus under elevated [CO₂] did not result in increased net N immobilization or C mineralization rates in long‐term laboratory incubations, possibly because microbial biomass was lower in the CO₂‐enriched plots. Furthermore, the δ¹³CO₂ of the C mineralized from the incubated soil closely tracked the δ¹³C of the labile POM pool in the elevated [CO₂] treatment, especially in shallower soil, and did not indicate significant priming of the decomposition of pre‐experiment soil organic matter (SOM). Although potential C mineralization rates were positively and linearly related to total SOM C content in the top 30 cm of soil, this relationship did not hold in deeper soil. Taken together with an increased mean residence time of C in deeper soil pools, these findings indicate that C inputs from relatively deep roots under elevated [CO₂] may increase the potential for long‐term soil C storage. However, C in deeper soil is likely to take many years to accrue to a significant fraction of total soil C given relatively smaller root inputs at depth. Expanded representation of biogeochemical cycling throughout the soil profile may improve model projections of future forest responses to rising atmospheric [CO₂].