重庆石灰岩地区主要木本植物叶片性状及养分再吸收特征

以重庆石灰岩地区15种常绿木本植物和14种落叶木本植物为研究对象,对两种生活型植物叶片衰老前后叶干物质含量(LDMC)、比叶面积(SLA)和叶片厚度(LT)进行了比较,并采用不同的计算方法(单位质量叶片养分含量、单位面积叶片养分含量)分析了两类植物叶片衰老前后养分含量及再吸收特征,最后对养分再吸收效率与其他叶性状因子之间的关系进行了相关分析。结果表明:常绿植物成熟叶LDMC、LT及衰老叶LT显著低于落叶植物,落叶植物成熟叶和衰老叶SLA均显著高于常绿植物(P0.05);基于单位质量叶片计算的养分含量,常绿植物成熟和衰老叶N、P量均低于落叶植物,而基于单位面积叶片计算的N、P含量则表现出相反的趋势;基于不同方法计算的N、P再吸收效率差异不明显,其中常绿植物基于单位质量叶片养分含量计算的N、P平均再吸收效率为39.42%、43.79%,落叶植物的为24.08%、33.59%;常绿和落叶植物N、P再吸收效率与LDMC、SLA、LT和成熟叶N、P含量之间没有显著相关性,但与衰老叶养分含量存在显著负相关(P0.05)。研究发现,无论是常绿植物还是落叶植物,衰老叶N、P含量均较低,表明石灰岩地区植物具有较高的养分再吸收程度。     

中亚热带植被恢复阶段植物叶片、凋落物、土壤碳氮磷化学计量特征

为揭示植被恢复过程中生态系统的养分循环机制及植物的生存策略, 根据亚热带森林群落演替过程, 采用空间代替时间方法, 以湘中丘陵区地域相邻、环境条件基本一致的檵木(Loropetalum chinensis) +南烛(Vaccinium bracteatu) +杜鹃(Rhododendron mariesii)灌草丛(LVR)、檵木+杉木(Cunninghamia lanceolata) +白栎(Quercus fabri)灌木林(LCQ)、马尾松(Pinus massoniana) +柯(Lithocarpus glaber) +檵木针阔混交林(PLL)、柯+红淡比(Cleyera japonica) +青冈(Cyclobalanopsis Glauca)常绿阔叶林(LCC)作为一个恢复系列, 设置固定样地, 采集植物叶片、未分解层凋落物和0-30 cm土壤样品, 测定有机碳(C)、全氮(N)、全磷(P)含量及其化学计量比, 运用异速生长关系、养分利用效率和再吸收效率分析植物对环境变化的响应和养分利用策略。结果表明: (1)随着植被恢复, 叶片C:N、C:P、N:P显著下降, 而叶片C、N、P含量和土壤C、N含量、C:P、N:P显著增加, 其中LCC植物叶片C、N含量, 土壤C、N含量及其N:P, PLL植物叶片P含量, 土壤C:P显著高于其他3个恢复阶段, 各恢复阶段植物叶片N:P > 20, 植物生长受P限制; 凋落物C、N、P含量及其化学计量比波动较大。(2)凋落物与叶片、土壤的化学计量特征之间的相关关系较弱, 叶片与土壤的化学计量特征之间具有显著相关关系, 其中叶片C、N、P含量与土壤C、N含量、C:N (除叶片C、N含量外)、C:P、N:P呈显著正相关关系; 叶片C:N与土壤C、N含量、C:P、N:P, 叶片C:P与土壤C含量、C:N、C:P, 叶片N:P与土壤C:N呈显著负相关关系。(3)植被恢复过程中, 叶片N、P之间具有显著异速生长关系, 异速生长指数为1.45, 叶片N、P的利用效率下降, 对N、P的再吸收效率增加, LCC叶片N利用效率最低, PLL叶片P利用效率最低而N、P再吸收效率最高。(4)叶片N含量内稳态弱, 而P含量具有较高的内稳态, 在土壤低P限制下植物能保持P平衡。植被恢复显著影响叶片、凋落物、土壤C、N、P含量及其化学计量比, 叶片与土壤之间C、N、P含量及化学计量比呈显著相关关系, 植物通过降低养分利用效率和提高养分再吸收效率适应土壤养分的变化, 叶片-凋落物-土壤系统的N、P循环随着植被恢复逐渐达到“化学计量平衡”。     

辽东山区主要阔叶树种叶片养分含量和再吸收对落叶时间的影响

凋落物是森林生态系统养分的重要来源, 叶片脱落时间是影响其分解的关键因素。东北温带森林中蒙古栎(Quercus mongolica)落叶时间较其他树种晚, 在山脊等贫瘠立地叶片甚至第二年春天才脱落。我们假设: 相对于其他树种, 蒙古栎叶片养分元素含量过高、再吸收时间长, 导致叶片延迟脱落。为验证假设, 除蒙古栎外, 选择了落叶时间居中的色木槭(Acer mono)和落叶较早的胡桃楸(Juglans mandshurica)为对象, 持续监测叶片从成熟至凋落过程中叶片养分元素含量, 包括大量元素: 氮(N)、磷(P)、钾(K)、钙(Ca)和镁(Mg), 微量元素: 铁(Fe)、铜(Cu)、锰(Mn)和锌(Zn); 并分析养分再吸收率。结果表明: 蒙古栎成熟叶养分元素含量介于对照树种之间; 凋落叶N、P和K含量低于对照树种, Fe和Mn含量高于对照树种, 其余元素含量介于对照树种之间。该结果不支持“蒙古栎叶片养分含量过高”假设。蒙古栎叶片N、P和K再吸收率高于对照树种, 再吸收率高低与其落叶时间完全一致; 叶片Cu和Zn再吸收率与对照树种无显著差异; 叶片其余元素未发生再吸收, 其累积率与对照树种无显著差异; 说明养分再吸收与养分含量无关, 可能与树种的种专一性相关, 可能会影响叶片脱落时间。由于蒙古栎多生长在贫瘠土壤, 其成熟叶无法积累更多养分; 为避免叶片脱落后养分进入土壤被其他物种利用, 将养分尽量回收储存于自身, 即蒙古栎叶片养分再吸收过程较长, 叶片脱落较晚。生长在极端贫瘠立地的蒙古栎叶片次年春天才落叶, 可能是由于再吸收一直在进行, 来不及脱落而保留至新生长季开始。落叶晚的树种养分再吸收率高、有利于自身养分保存, 更能适应贫瘠土壤, 反之亦然。     

Effects of leaf nutrient concentration and resorption on leaf falling time of dominant broadleaved species in a montane region of eastern Liaoning Province,China

凋落物是森林生态系统养分的重要来源, 叶片脱落时间是影响其分解的关键因素。东北温带森林中蒙古栎(Quercus mongolica)落叶时间较其他树种晚, 在山脊等贫瘠立地叶片甚至第二年春天才脱落。我们假设: 相对于其他树种, 蒙古栎叶片养分元素含量过高、再吸收时间长, 导致叶片延迟脱落。为验证假设, 除蒙古栎外, 选择了落叶时间居中的色木槭(Acer mono)和落叶较早的胡桃楸(Juglans mandshurica)为对象, 持续监测叶片从成熟至凋落过程中叶片养分元素含量, 包括大量元素: 氮(N)、磷(P)、钾(K)、钙(Ca)和镁(Mg), 微量元素: 铁(Fe)、铜(Cu)、锰(Mn)和锌(Zn); 并分析养分再吸收率。结果表明: 蒙古栎成熟叶养分元素含量介于对照树种之间; 凋落叶N、P和K含量低于对照树种, Fe和Mn含量高于对照树种, 其余元素含量介于对照树种之间。该结果不支持“蒙古栎叶片养分含量过高”假设。蒙古栎叶片N、P和K再吸收率高于对照树种, 再吸收率高低与其落叶时间完全一致; 叶片Cu和Zn再吸收率与对照树种无显著差异; 叶片其余元素未发生再吸收, 其累积率与对照树种无显著差异; 说明养分再吸收与养分含量无关, 可能与树种的种专一性相关, 可能会影响叶片脱落时间。由于蒙古栎多生长在贫瘠土壤, 其成熟叶无法积累更多养分; 为避免叶片脱落后养分进入土壤被其他物种利用, 将养分尽量回收储存于自身, 即蒙古栎叶片养分再吸收过程较长, 叶片脱落较晚。生长在极端贫瘠立地的蒙古栎叶片次年春天才落叶, 可能是由于再吸收一直在进行, 来不及脱落而保留至新生长季开始。落叶晚的树种养分再吸收率高、有利于自身养分保存, 更能适应贫瘠土壤, 反之亦然。     

两种林龄文冠果叶N、P、K的季节变化及再吸收特征

以科尔沁沙地西南缘乌丹地区文冠果人工林为研究对象,分析了8年生和50年生文冠果叶片N、P、K含量的季节变化和再吸收效率。结果表明:2种林龄文冠果年生长周期内N、P、K含量均呈下降趋势,其中,5—6月和8—10月下降幅度较大,6—8月含量变化不大;8年生文冠果叶片凋落前后的N、P、K含量(衰落叶中N除外)均极显著50年生文冠果(P0.01)。8年生文冠果叶片养分再吸收效率表现为:N(50%)P(32%)K(30%),但50年生的则表现为K(63%)P(62%)N(48%);二者N再吸收效率差异不显著(P0.05),而50年生文冠果叶片P、K再吸收效率极显著高于8年生文冠果(P0.01)。50年生文冠果叶片再吸收效率高,反映其较高的养分保存能力和养分利用效率,能更好地适应贫瘠养分生境。     

长白山阔叶红松林5种主要树种水分利用效率与叶片养分特征

以长白山阔叶红松林中5种优势树种(红松、水曲柳、色木槭、蒙古栎、紫椴)为研究对象,利用稳定碳同位素技术分析了植物的水分利用效率(WUE),测定了叶片碳(C)、氮(N)、磷(P)3种营养元素的含量用以分析植物对养分的利用特征,并综合分析了植物WUE与叶片养分含量之间的关系.结果 表明:冠层不同位置微气象因子的差异导致WU...     

地表连年覆盖雷竹林叶片养分利用特征

对连续覆盖雷竹林换叶期林地土壤和叶片养分利用效率、养分再吸收效率进行了研究,旨在阐明地表连年覆盖栽培对雷竹林养分利用特征的影响.结果表明:连续覆盖3年雷竹林土壤N养分有效性、N素利用效率以及再吸收效率均显著性提高,雷竹林表现出较高的保存N养分的能力;土壤P元素有效性水平显著升高,而叶片P养分的利用效率显著降低,反映出叶片对土壤高P含量的适应策略,N、P养分利用效率之间存在显著的负相关关系(P＜0.05);土壤K元素的有效性水平显著降低,竹林通过提高自身的K再吸收效率来维持养分的正常循环,再吸收效率高达95.8％;不同覆盖年限雷竹林Nm/Pm均＜14,连续覆盖3年雷竹林Nm/Pm比值极显著下降,经推断是此时雷竹林对土壤有效P高吸收的结果.成熟叶片的Nm/Pm与N、P的再吸收效率均无显著相关性,Nm/Pm并不能很好地反映叶片在凋落前的养分再分配格局.     

中亚热带植被恢复阶段植物叶片、凋落物、土壤碳氮磷化学计量特征

为揭示植被恢复过程中生态系统的养分循环机制及植物的生存策略,根据亚热带森林群落演替过程,采用空间代替时间方法,以湘中丘陵区地域相邻、环境条件基本一致的檵木(Loropetalumchinensis)+南烛(Vacciniumbracteatu)+杜鹃(Rhododendron mariesii)灌草丛(LVR)、檵木+杉木(Cunninghamia lanceolata)+白栎(Quercus fabri)灌木林(LCQ)、马尾松(Pinus massoniana)+柯(Lithocarpus glaber)+檵木针阔混交林(PLL)、柯+红淡比(Cleyera japonica)+青冈(Cyclobalanopsis Glauca)常绿阔叶林(LCC)作为一个恢复系列,设置固定样地,采集植物叶片、未分解层凋落物和0–30 cm土壤样品,测定有机碳(C)、全氮(N)、全磷(P)含量及其化学计量比,运用异速生长关系、养分利用效率和再吸收效率分析植物对环境变化的响应和养分利用策略。结果表明:(1)随着植被恢复,叶片C:N、C:P、N:P显著下降,而叶片C、N、P含量和土壤C、N含量、C:P、N:P显著增加,其中LCC植物叶片C、N含量,土壤C、N含量及其N:P,PLL植物叶片P含量,土壤C:P显著高于其他3个恢复阶段,各恢复阶段植物叶片N:P 20,植物生长受P限制;凋落物C、N、P含量及其化学计量比波动较大。(2)凋落物与叶片、土壤的化学计量特征之间的相关关系较弱,叶片与土壤的化学计量特征之间具有显著相关关系,其中叶片C、N、P含量与土壤C、N含量、C:N (除叶片C、N含量外)、C:P、N:P呈显著正相关关系;叶片C:N与土壤C、N含量、C:P、N:P,叶片C:P与土壤C含量、C:N、C:P,叶片N:P与土壤C:N呈显著负相关关系。(3)植被恢复过程中,叶片N、P之间具有显著异速生长关系,异速生长指数为1.45,叶片N、P的利用效率下降,对N、P的再吸收效率增加, LCC叶片N利用效率最低, PLL叶片P利用效率最低而N、P再吸收效率最高。(4)叶片N含量内稳态弱,而P含量具有较高的内稳态,在土壤低P限制下植物能保持P平衡。植被恢复显著影响叶片、凋落物、土壤C、N、P含量及其化学计量比,叶片与土壤之间C、N、P含量及化学计量比呈显著相关关系,植物通过降低养分利用效率和提高养分再吸收效率适应土壤养分的变化,叶片-凋落物-土壤系统的N、P循环随着植被恢复逐渐达到"化学计量平衡"。     

皖南亚热带常绿阔叶林林下灌木层主要树种叶片养分特征

对皖南亚热带常绿阔叶林林下灌木层主要树种草珊瑚〔Sarcandra glabra(Thunb.)Nakai〕、乌药〔Lindera aggregata(Sims)Kosterm.〕、香桂(Cinnamomum subavenium Miq.)、红楠(Machilus thunbergii Sieb.et Zucc.)、连蕊茶〔Camellia cuspidata(Kochs)H.J.Veitch〕、甜槠〔Castanopsis eyrei(Champ.ex Benth.)Tutch.〕、矩叶鼠刺(Itea oblonga Hand.-Mazz.)、红淡比(Cleyera japonica Thunb.)和小叶蚊母树〔Distylium buxifolium(Hance)Merr.〕叶片养分(包括C、N、P、K、Ca和Mg)含量、单位叶面积养分含量和比叶面积进行了测定,并分析了比叶面积与养分含量和单位叶面积养分含量的相关性。结果表明:9种树种叶片的C、N、P、K、Ca和Mg含量分别为438.09~492.31、9.79~15.60、0.33~2.06、11.39~32.52、3.84~13.34和2.85~14.05 g·kg-1,单位叶面积C、N、P、K、Ca和Mg含量分别为24.48~47.38、0.66~1.03、0.02~0.21、0.73~1.82、0.22~1.35和0.20~1.05 g·m-2,C/N比、C/P比和N/P比分别为31.40~47.88、227.76~1 495.66和5.26~32.90。9种树种叶片的比叶面积平均值为14.5 m2·kg-1,其中草珊瑚的比叶面积最大(17.90 m2·kg-1)、小叶蚊母树的比叶面积最小(9.89 m2·kg-1)。相关性分析结果表明:供试树种叶片的C/P比与N/P比间呈极显著正相关;比叶面积与叶片N含量呈显著正相关,与单位叶面积C、N和Ca含量分别呈极显著和显著负相关,与其他指标的相关性均不显著。综合分析结果表明:皖南亚热带常绿阔叶林下灌木层主要树种叶片的养分含量及比叶面积差异明显,其中,乌药、草珊瑚、矩叶鼠刺和香桂叶片的养分含量较高,而小叶蚊母树、红淡比和红楠的单位叶面积养分含量较高。     

密度对四季竹叶片C、N、P化学计量和养分重吸收特征的影响

以四季竹纯林为对象,研究了4种密度(D1:24600～29800株·hm-2;D2:37500～42600株·hm-2;D3:46500～52800株·hm-2;D4:76500～85500株·hm-2)下1～3年生立竹的成熟叶和凋萎叶的C、N、P化学计量特征及养分重吸收规律.结果表明:随竹林密度的增大,成熟叶C、N、P含量和凋萎叶C、P含量总体下降,凋萎叶N含量先升高后降低,D4密度时成熟叶C含量急剧下降;叶片C/N和C/P呈升高趋势,N/P先升高后降低.D3、D4密度的叶片N、P养分利用效率显著高于D1、D2密度;叶片N、P重吸收率分别呈先降后升和持续升高的趋势.D1～D3密度的成熟叶N/P为16.24～19.37,四季竹生长受P的限制,促进叶片建成,种群生长旺盛.D4密度的成熟叶N/P为13.42～15.74,N限制作用增强,叶片凋萎脱落,种群生长受到抑制.四季竹通过调节叶片的C、N、P含量及其化学计量比,提高了N、P利用效率及重吸收能力,以适应高密度下环境资源的激烈竞争.在本试验条件下,D3密度是四季竹适宜的经营密度.     

Photosynthesis-Nitrogen Relationships in Pioneer Plants of Disturbed Tropical Montane Forest Sites

Tropical forest disturbances lead to the establishment of secondary successional plant communities constituted by light demanding species with high relative growth rate that conduct to rapid canopy closure. Two main strategies for N nutrition are: (a) mineral N acquisition in the form of NH4 and NO3, and (b) symbiotic atmospheric N2 fixation. Given the high N requirement for maximization of leaf area and radiant energy absorption, we hypothesize that contrasting strategies of N nutrition in these environments are reflected in leaf photosynthetic characteristics. We compared the N-photosynthesis relationships and carbon balance parameters per unit leaf area as they vary with age in two species with contrasting N acquisition strategies: a N2-fixer Crotalaria anagyroides HBK (Papilionoideae), and a mineral-N user Verbesina turbacensis HBK (Asteraceae). N2 fixation capacity was associated to higher specific leaf area (SLA), higher photosynthetic capacity (Pmax) per unit leaf area and leaf mass, and higher N content per unit leaf mass. The N2-fixer species showed higher slope in the relationship Pmax-N per unit leaf mass and area when compared to the leaves of non-fixer species. Moreover, the intrinsic photosynthetic N use efficiency (Pmax/N) was higher in the N2 fixer than in leaves of the non-fixer species. Changes in N due to leaf age resulted in larger changes in CO2 flux density at the leaf level in the N2-fixer species. The higher photosynthetic capacity of the N2-fixer species was mechanistically related to higher stomatal conductance, internal CO2 concentration (ci) values closer to atmospheric CO2 concentration (ca), and lower intrinsic water use efficiency than the mineral N-user species. Despite their higher Pmax per unit leaf area, total non-structural saccharides concentration was lower in mature leaves of the N2-fixer plant as compared to the non-fixer counterpart. This might be caused by the presence of a larger root sink (symbionts) stimulating saccharides export and higher diurnal respiration rates.     

Mean residence time of leaf number, area, mass, and nitrogen in canopy photosynthesis

Mean residence time (MRT) of plant nitrogen (N), which is an indicator of the expected length of time N newly taken up is retained before being lost, is an important component in plant nitrogen use. Here we extend the concept MRT to cover such variables as leaf number, leaf area, leaf dry mass, and nitrogen in the canopy. MRT was calculated from leaf duration (i.e., time integral of standing amount) divided by the total production of leaf variables. We determined MRT in a Xanthium canadense stand established with high or low N availability. The MRT of leaf number may imply longevity of leaves in the canopy. We found that the MRT of leaf area and dry mass were shorter than that of leaf number, while the MRT of leaf N was longer. The relatively longer MRT of leaf N was due to N resorption before leaf shedding. The MRT of all variables was longer at low N availability. Leaf productivity is the rate of canopy photosynthesis per unit amount of leaf variables, and multiplication of leaf productivity by MRT gives the leaf photosynthetic efficiency (canopy photosynthesis per unit production of leaf variables). The photosynthetic efficiency of leaf number implies the lifetime carbon gain of a leaf in the canopy. The analysis of plant-level N use efficiency by evaluating the N productivity and MRT is a well-established approach. Extension of these concepts to leaf number, area, mass, and N in the canopy will clarify the underlying logic in the study of leaf life span, leaf area development, and dry mass and N use in canopy photosynthesis.     

Canopy structure and vertical patterns of photosynthesis and related leaf traits in a deciduous forest

Canopy structure and light interception were measured in an 18-m tall, closed canopy deciduous forest of sugar maple (Acer saccharum) in southwestern Wisconsin, USA, and related to leaf structural characteristics, N content, and leaf photosynthetic capacity. Light attenuation in the forest occurred primarily in the upper and middle portions of the canopy. Forest stand leaf area index (LAI) and its distribution with respect to canopy height were estimated from canopy transmittance values independently verified with a combined leaf litterfall and point-intersect method. Leaf mass, N and A _max per unit area (LMA, N/area and A _max/area, respectively) all decreased continuously by over two-fold from the upper to lower canopy, and these traits were strongly correlated with cumulative leaf area above the leaf position in the canopy. In contrast, neither N concentration nor A _max per unit mass varied significantly in relation to the vertical canopy gradient. Since leaf N concentration showed no consistent pattern with respect to canopy position, the observed vertical pattern in N/area is a direct consequence of vertical variation of LMA. N/area and LMA were strongly correlated with A _max/area among different canopy positions (r²=0.81 and r²=0.66, respectively), indicating that vertical variation in area-based photosynthetic capacity can also be attributed to variation in LMA. A model of whole-canopy photosynthesis was used to show that observed or hypothetical canopy mass distributions toward higher LMA (and hence higher N/area) in the upper portions of the canopy tended to increase integrated daily canopy photosynthesis over other LMA distribution patterns. Empirical relationships between leaf and canopy-level characteristics may help resolve problems associated with scaling gas exchange measurements made at the leaf level to the individual tree crown and forest canopy-level.     

Nutrient content in Quercus ilex canopies: Seasonal and spatial variation within a catchment

Spatial and temporal changes in canopy nutrient content were studied in 1988 and 1989 in a Mediterranean Quercus ilex ssp. ilex L. forest in north-eastern Spain. Sampling was conducted in parallel at two sites which represent endpoints along a slope gradient within a small catchment (ridge top at 975 m and valley bottom at 700 m). Deeper soils resulted in significantly higher N and P concentrations, and N content on a leaf area basis at the valley bottom site. In contrast, K concentration in leaves was significantly higher at the ridge top site, where soil K concentration was also higher. At both sites, N and P content on a leaf area basis was highest at the top of the canopy, where leaf area is highest. N resorption efficiency decreased from top to bottom of the canopy. Results suggested a minor role of shaded leaves as nutrient storage sites. Lower P resorption efficiency was found at the ridge top site. Seasonal changes of P and N concentration on a leaf area basis suggest P replenishment, and to a lesser degree N, during periods of lower growth activity due to low temperatures, but coinciding with higher water availability (autumn-early spring period). Thus, N and P resorption from the remaining foliage in the canopy took place, and to a larger degree at the valley bottom site, coinciding with a slightly higher leaf area index and productivity at this site.     

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     

A leaf gas exchange model that accounts for intra-canopy variability by considering leaf nitrogen content and local acclimation to radiation in grapevine (Vitis vinifera L.)

Understanding the distribution of gas exchange within a plant is a prerequisite for scaling up from leaves to canopies. We evaluated whether leaf traits were reliable predictors of the effects of leaf ageing and leaf irradiance on leaf photosynthetic capacity (V(cmax) , J(max) ) in field-grown vines (Vitis vinifera L). Simultaneously, we measured gas exchange, leaf mass per area (LMA) and nitrogen content (N(m) ) of leaves at different positions within the canopy and at different phenological stages. Daily mean leaf irradiance cumulated over 10 d (PPFD(10) ) was obtained by 3D modelling of the canopy structure. N(m) decreased over the season in parallel to leaf ageing while LMA was mainly affected by leaf position. PPFD(10) explained 66, 28 and 73% of the variation of LMA, N(m) and nitrogen content per area (N(a) ), respectively. Nitrogen content per unit area (N(a) = LMA × N(m) ) was the best predictor of the intra-canopy variability of leaf photosynthetic capacity. Finally, we developed a classical photosynthesis-stomatal conductance submodel and by introducing N(a) as an input, the model accurately simulated the daily pattern of gas exchange for leaves at different positions in the canopy and at different phenological stages during the season.     

Nitrogen fertilization enhances water‐use efficiency in a saline environment

Effects of salinity and nutrients on carbon gain in relation to water use were studied in the grey mangrove, Avicennia marina, growing along a natural salinity gradient in south‐eastern Australia. Tall trees characterized areas of seawater salinities (fringe zone) and stunted trees dominated landward hypersaline areas (scrub zone). Trees were fertilized with nitrogen (+N) or phosphorus (+P) or unfertilized. There was no significant effect of +P on shoot growth, whereas +N enhanced canopy development, particularly in scrub trees. Scrub trees maintained greater CO₂ assimilation per unit water transpired (water‐use efficiency, WUE) and had lower nitrogen‐use efficiency (NUE; CO₂ assimilation rate per unit leaf nitrogen) than fringe trees. The CO₂ assimilation rates of +N trees were similar to those in other treatments, but were achieved at lower transpiration rates, stomatal conductance and intercellular CO₂ concentrations. Maintaining comparable assimilation rates at lower stomatal conductance requires greater ribulose 1·5‐bisphosphate carboxylase/oxygenase activity, consistent with greater N content per unit leaf area in +N trees. Hence, +N enhanced WUE at the expense of NUE. Instantaneous WUE estimates were supported by less negative foliar δ¹³C values for +N trees and scrub control trees. Thus, nutrient enrichment may alter the structure and function of mangrove forests along salinity gradients.     

The influence of phosphorus deficiency on growth and nitrogen fixation of white clover plants

The effects of P deficiency on growth, N(2)-fixation and photosynthesis in white clover (Trifolium repens L.) plants were investigated using three contrasting relative addition rates of P, or following abrupt withdrawal of the P supply. Responses to a constant below-optimum P supply rate consisted of a decline in N(2)-fixation per unit root weight and a small reduction in the efficiency with which electrons were allocated to the reduction of N(2) in nodules. Abrupt removal of P arrested nodule growth and caused a substantial decline in nitrogenase activity per unit root weight, but not per unit nodule mass. Similarly, the rate of photosynthesis per unit leaf area was unaffected by abrupt P removal, whereas CO(2) acquisition for the plant as a whole decreased due to a decline in total leaf area, leaf area per unit leaf weight and utilization of incoming radiation. These changes followed the decline in tissue P concentrations. The ratio between CO(2)-fixation and N(2)-fixation was maintained under short-term P deprivation but increased under long-term low P supply, indicating a regulatory inhibition of nodule activity following morphological and growth adjustments. It is concluded that N(2)-fixation did not limit the growth of clover plants experiencing P deficiency. A low P status induced changes in the relative growth of roots, nodules and shoots rather than changes in N and/or C uptake rates per unit mass or area of these organs.     

Forest growth along a rainfall gradient in Hawaii: Acacia koa stand structure,productivity, foliar nutrients,and water- and nutrient-use efficiencies

We tested whether variation in growth of native koa (Acacia koa) forest along a rainfall gradient was attributable to differences in leaf area index (LAI) or to differences in physiological performance per unit of leaf area. Koa stands were studied on western Kauai prior to Hurricane Iniki, and ranged from 500 to 1130 m elevation and from 850 to 1800 mm annual precipitation. Koa stands along the gradient had basal area ranging from 8 to 42 m²/ha, LAI ranging from 1.4 to 5.4, and wood increment ranging from 0.7 to 7.1 tonnes/ha/year. N, P, and K contents by weight of sun leaves (phyllodes) were negatively correlated with specific leaf mass (SLM, g m^-2) across sites; on a leaf area basis, N increased whereas P and K decreased with SLM. LAI, aboveground woody biomass increment, and production per unit leaf area (E) increased as phyllode ¹³C became more negative. The ¹³C data suggested that intrinsic water-use efficiency (ratio of assimilation to conductance) increased as water availability decreased. In five of the six sites, phyllode P contents increased as LAI increased, but biomass increment and E were not correlated with phyllode nutrient contents, suggesting that productivity was limited more by water than by nutrient availability. Because vapor pressure deficits increased with decreasing elevation, actual water-use efficiency (ratio of assimilation to transpiration) was lower at drier, low-elevation sites. There was a trade-off between intrinsic water-use efficiency and production per unit of canopy N or P across the gradient. In summary, koa responds to water limitation both by reducing stand LAI and by adjusting gas exchange, which results in increased intrinsic water-use efficiency but decreased E.