黄土高原典型小流域草地群落功能性状对土壤水分的响应

植物功能性状可以响应生境的变化并决定生态系统的功能,探究植物功能性状间的关系及其随土壤有效水分梯度的变化规律,对认识不同水分条件下植被在群落水平碳水代谢关系和维持水分平衡的生理生态学机制具有重要意义。以甘肃定西典型半干旱黄土小流域草地群落为研究对象,采用排序分析和回归拟合方法,分析了30个代表性草地样地中7个植物功能性状加权均值对土壤有效水分的响应以及响应性状间的相关关系。结果显示:(1)7个性状中,除叶宽与土壤有效水分无明显相关外,叶长、株高、叶面积、比叶面积、叶厚和叶干物质含量均与土壤有效水分显著性相关,可识别为草地在群落水平对土壤水分的响应性状。(2)草地群落通过降低株高,减小叶长、叶面积和比叶面积,增加叶厚和叶干物质含量以适应土壤有效水分减少;其中叶干物质含量的解释度最大,是土壤水分的最优响应性状。(3)除叶厚与叶长无显著相关外,其余功能性状均存在显著相关,说明草地群落的功能性状在土壤水分梯度上已基本形成了一个相互权衡或协同变化的功能性状组合。     

武夷山不同海拔黄山松枝叶大小关系

对武夷山自然保护区不同海拔黄山松叶片面积、叶片数量、小枝长度及小枝直径等性状进行测定,分析不同海拔黄山松枝-叶大小间的权衡关系.结果表明: 随海拔升高,黄山松小枝的叶片数量、小枝长度、小枝直径、出叶强度及茎截面积逐渐增大,单叶面积呈逐渐减小趋势;不同海拔黄山松小枝出叶强度与单叶面积均呈显著负相关,不同海拔黄山松小枝茎截面积与总叶面积呈显著正相关;不同海拔黄山松小枝长度、小枝直径与出叶强度呈显著负相关,与单叶面积、叶片数量及总叶面积均呈显著正相关.为提高竞争优势或是资源利用效率,低海拔黄山松倾向于在短枝上着生量少但单叶面积大的针叶,而高海拔黄山松趋向于在长枝上着生量大但单叶面积小的针叶,这体现出不同海拔梯度黄山松小枝的资源利用策略及枝叶间生物量分配的权衡机制.     

黔中喀斯特木本植物功能性状变异及其适应策略

性状变异反映了植物的生活史对策。该研究以贵州普定县天龙山10种木本植物为对象, 通过分析枝叶和根系9个功能性状的种间与种内变异, 揭示植物对喀斯特生境的适应策略。结果表明: (1) 9个性状变异程度不同, 细根组织密度的种间和种内变异系数最大, 分别达96.47%和51.44%, 小枝干物质含量的种间与种内变异最小, 分别为11.67%和6.83%。(2)种间水平的细根组织密度在不同物种中没有显著的差异, 比根长、叶厚度、叶面积、比叶面积、叶干物质含量、叶组织密度、小枝干物质含量和小枝组织密度均表现出显著的差异。在种内, 比叶面积差异显著, 其他性状差异不显著。(3)绝大多数叶和枝性状间显著相关, 比根长与比叶面积显著负相关, 其他根系性状与枝叶性状相关性不显著。总之, 与同纬度非喀斯特地区植物相比, 普定喀斯特地区植物具有较小的叶面积和比根长度, 较大的叶干物质含量、叶组织密度等一系列有利于减小蒸腾和储存养分的功能性状组合, 这可能是其适应干旱贫瘠的喀斯特环境的主要生态策略。     

胡杨当年生小枝茎构型对展叶效率的影响

枝叶异速生长关系对理解荒漠植物生物量分配和生活史策略具有重要意义。该研究以小枝茎长度、茎纤细率和茎体积表征茎构型, 以叶密度(单位茎长度的叶数量)、叶面积比(单位小枝茎干质量的总叶面积)和叶茎质量比(单位小枝茎干质量的总叶干质量)表征展叶效率, 采用标准化主轴回归(SMA)方法研究胡杨(Populus euphratica)当年生小枝茎构型对展叶效率的影响及二者沿地下水埋深(GWD)梯度的权衡策略。结果显示, 胡杨当年生小枝茎直径、展叶效率和比叶面积、叶大小随GWD增加而降低, 小枝茎长度、茎纤细率及其上的叶数量则增大。小枝茎构型性状均与展叶效率呈显著负相关关系, 即随小枝茎长度、茎纤细率和茎体积的增加, 展叶效率逐渐降低, 这可能是枝叶大小、水分传导与机械支撑间的权衡结果。小枝茎构型与展叶效率的异速生长指数(斜率)随GWD增加而增大, 是由于单位小枝茎投资获得的叶面积或者叶质量降低所致, 反映出随GWD增加胡杨采取了高消耗低收益的保守型策略。胡杨应对环境压力时, 倾向于在长枝上着生数量较多的小叶, 短枝上着生数量较少的大叶, 体现出胡杨小枝的资源利用策略与枝叶大小的权衡机制。综上所述, GWD显著影响胡杨小枝茎构型-展叶效率的权衡关系, 低展叶效率是胡杨应对日益旱化荒漠环境的适应策略。     

Variations of plant functional traits and adaptive strategy of woody species in a karst forest of central Guizhou Province,southwestern China

性状变异反映了植物的生活史对策。该研究以贵州普定县天龙山10种木本植物为对象, 通过分析枝叶和根系9个功能性状的种间与种内变异, 揭示植物对喀斯特生境的适应策略。结果表明: (1) 9个性状变异程度不同, 细根组织密度的种间和种内变异系数最大, 分别达96.47%和51.44%, 小枝干物质含量的种间与种内变异最小, 分别为11.67%和6.83%。(2)种间水平的细根组织密度在不同物种中没有显著的差异, 比根长、叶厚度、叶面积、比叶面积、叶干物质含量、叶组织密度、小枝干物质含量和小枝组织密度均表现出显著的差异。在种内, 比叶面积差异显著, 其他性状差异不显著。(3)绝大多数叶和枝性状间显著相关, 比根长与比叶面积显著负相关, 其他根系性状与枝叶性状相关性不显著。总之, 与同纬度非喀斯特地区植物相比, 普定喀斯特地区植物具有较小的叶面积和比根长度, 较大的叶干物质含量、叶组织密度等一系列有利于减小蒸腾和储存养分的功能性状组合, 这可能是其适应干旱贫瘠的喀斯特环境的主要生态策略。     

阔叶红松林6种阔叶树单个叶片叶面积和叶干质量的垂直变异及模型构建

快捷、准确地测定植物叶片尺度上的叶面积(LA)和叶干质量(LDM)对于探讨植物性状对气候变化的响应机制至关重要,但适于测定区域尺度上不同阔叶植物单个叶片LA和LDM的经验模型尚未提出.本研究以中国东北4个分布区阔叶红松林内的白桦、紫椴、山杨、枫桦、水曲柳和裂叶榆6种阔叶树种为研究对象,分别测定其不同冠层高度(上层、中层和下层)叶片的叶长、叶宽、叶厚、LA及LDM.以叶长与叶宽之比(叶长宽比)的中位数为标准将6种阔叶树分为两组,检验每组树种不同冠层高度对构建预测LA和LDM的经验模型是否存在显著影响;构建适于预测区域尺度上不同冠层单个叶片LA和LDM的经验模型,验证其预测精度;并进一步评估构建的经验模型预测其他区域相同阔叶树种LA和LDM的适用性.结果表明: 整体上6种阔叶树单个叶片的LA随冠层高度的降低呈显著增大趋势,而部分树种的LDM呈下降趋势;冠层高度对构建预测LA和LDM的经验模型具有显著影响;构建的经验模型预测两组阔叶树种不同冠层单个叶片LA和LDM的平均精度分别为95%和83%,且基于这些模型预测其他区域相应树种LA和LDM的平均精度分别为94%和80%,表明本研究构建的经验模型在我国东北区域具有普遍适用性.     

同期生长在种源地与异地同质园刨花楠叶性状分析

为辨别环境变化与遗传因素对植物叶片主要功能性状的影响,以同期生长在4种源地母树林下及异地同质园的1.5年生刨花楠苗木为研究对象,对其叶片表型及养分性状进行对比分析。结果显示：(1)刨花楠叶面积、叶厚、叶干物质含量等叶片表型性状受遗传与环境因素共同影响;叶片碳(C)含量受遗传因素调控,环境对其影响较小;叶片氮、磷(N、P)含量主要受环境因素影响;(2)不同种源刨花楠比叶面积、叶厚、叶干物质含量、叶形指数等性状变异系数较大(8.85%—37.03%),其中江西遂川种源变异系数相对较大,而湖南茶陵种源则相对较小,各种源都倾向于通过调节比叶面积、叶厚、叶片氮磷含量等性状以适应生境变化;(3)种源地与同质园刨花楠的比叶面积虽均与叶厚呈显著负相关,但同质园刨花楠比叶面积与叶形指数呈显著正相关,与叶片氮含量无明显相关,而种源地刨花楠比叶面积则与叶形指数无明显相关,与叶片氮含量则呈显著负相关;(4)不同种源苗木叶性状指标在种源地与同质园间存在不同的协调与权衡,体现了植物在不同生境下的适应策略。其中湖南茶陵种源在两种生境下都具有更保守的资源获取策略,而江西安福种源对环境变化则更为敏感,资源获取策略更为灵...     

浙江天童木本植物Corner法则的检验:个体密度的影响

Corner法则反映了植物枝叶大小和数量配置的构型策略,但是,对于个体密度如何影响枝叶关系的理解仍不够深入。该研究选择浙江天童的25个植物群落,通过比较枝大小(横截面积)-叶大小(总叶面积)关系和枝大小(横截面积)-枝数量(分梢密度)关系,分析个体竞争对植物Corner法则的影响。结果显示:1)在不同密度区间,枝横截面积和总叶面积均显著异速正相关。2)个体水平上,枝大小-叶大小回归方程的截距在低密度区间显著小于高密度区间,表明在枝大小一定的条件下,高密度群落的植物当年生枝条会支撑更大的总叶面积;而物种水平上,枝大小-叶大小回归方程的截距在不同密度区间没有显著性差异。3)枝横截面积与分梢密度显著负相关,且各密度区间也存在显著小于–1的共斜率。4)个体水平和物种水平的分析结果都显示,枝横截面积与分梢密度回归方程的截距在不同密度区间无显著差异,表明高密度植物并没有比低密度植物在单位大小枝条上配置更多的分枝。总之,植物枝大小-叶大小关系和枝大小-枝数量关系各自在不同的密度区间具有共同的变化斜率,反映了天童地区植物Corner法则不随个体密度变化而改变。但是,枝叶关系回归方程截距的改变表明,个体竞争的加大会使得植物在枝叶大小的配置策略上进行调整,从而可能通过生态位分化促进物种共存。     

薰衣草枝叶性状关系的个体大小依赖

植物枝叶性状的个体大小差异,是植物适应异质性环境所形成的冠层构建策略,对于理解枝叶构建机制及光合生理代谢具有重要意义。于2017年7月下旬,在金水湖湿地公园选择一块薰衣草样地,根据体积将薰衣草(Lavandula angustifolia)分为3个大小等级[I级:植株体积的立方根(d)≤60 cm、II级(60 cmd≤90 cm)和III级(d90 cm)],采用一元线性回归方法,研究了薰衣草种群枝叶性状的个体大小依赖。结果表明:随着薰衣草植株大小等级增大,薰衣草的叶面积、枝长度、枝数量和枝横截面积逐渐增大,而叶数量、叶厚度和分枝角度逐渐减小。薰衣草叶面积、枝长度和枝数量与个体大小呈极显著的正相关(P0.01),枝横截面积与个体大小呈显著的正相关(P0.05),叶数量和叶厚度与个体大小呈极显著的负相关(P0.01),分枝角度与个体大小呈显著的负相关(P0.01)。为提高资源利用效率,大个体薰衣草选择生长少量大而薄的叶片以及分配更多的生物量用于小枝的生长;而小个体薰衣草选择生长多数小而厚的叶片以及短而细的枝条,体现了不同大小等级薰衣草枝叶表型可塑性。     

浙江天童木本植物Corner法则的检验:微地形的影响

枝叶的大小和数量关系表征了植物应对环境胁迫的水力结构特征,当前,对其在微生境间的变化规律不太清楚。该研究在浙江天童常绿阔叶林中,按照地形的凸凹差异,各选择了10个群落,采用标准化主轴估计方法,分别比较了植物枝大小(横截面积)-叶大小(总叶面积)关系、枝大小(横截面积)-枝数量(枝稠密度)关系和叶大小(单叶面积)-叶数量(生叶强度)关系在两类生境的变化。结果发现:1)在两类生境中,枝横截面积和总叶面积显著异速正相关(p0.001),存在显著大于1的共斜率;其回归方程截距在凹型生境显著大于凸型生境,表明在一定枝大小下,凹型生境比凸型生境的植物枝条支撑更大的叶面积;2)枝横截面积与枝稠密度显著负相关(p0.001),且在两生境间存在显著小于–1的共斜率;回归方程截距在生境间无显著差异(p0.05),表明凹型与凸型生境的植物在单位大小枝条配置的分枝数相同;3)单叶面积与生叶强度显著负相关(p0.001),且也在生境间存在显著小于–1的共斜率;回归方程的截距在凹型生境显著大于凸型生境,表明在叶片大小相同时,凹型比凸型生境的植物在单位大小枝条上支撑的叶片数更多。综上,与凸型生境相比,凹型生境植物单位大小枝条更倾向于支持数量较多的大叶片。本研究证明,植物Corner法则和叶大小-叶数量的权衡在局域尺度也具普适性。枝叶大小和叶数量配置的调整体现了植物水力结构对凸凹生境不同水分供应的选择偏好。     

亚热带常绿阔叶林植物叶小枝的异速生长

 植物生态学研究的重要内容之一是识别和定量刻画种间生态变异的主要维数,叶大小小枝大小维（谱）是其中之一,目前的研究相对比较薄弱,两者之间是异速还是等速生长关系仍存在着争论。亚热带常绿阔叶林植物叶大小-枝大小维的研究报道很少。该文以我国东部亚热带典型区域福建梅花山常绿阔叶林的68种常绿乔灌木植物为对象,进行了叶-小枝关系及其生态意义的研究。结果表明：1）小枝茎截面积与叶干重、总叶面积和单叶面积之间的SMA斜率分别为1.29、1.23和1.18,呈现异速生长关系,支持叶大小 小枝大小为异速生长的相关研究结论,但SMA斜率低于预期值,其原因及生态意义有待进一步研究;2）小枝总叶面积与单叶面积呈显著正相关,而与叶片数量不相关,反映了小枝总叶面积的增加主要是由单叶面积大小决定的,可能与这一地区湿润气候有关;而单叶面积与枝条长度呈正相关则可能反映了植物对常绿阔叶林内较弱光照环境的适应;3）叶干重同小枝干重、叶面积为等速生长关系,可能反映了植物与动物之间代谢方式的差异。     

Effects of Wind on the Allometry of Two Species of Plants in an Elfin Cloud Forest

Thigmomorphogenesis includes the effects of mechanical perturbation on plant growth. To test whether thigmomorphogenesis is evident at different scales within plants, we investigated the effect of wind on allometric relationships between specific plant parts. We chose two species from the elfin cloud forest of Puerto Rico that have contrasting growth habits, the shrub Clibadiun erosum (Asteraceae) and the palm Prestoea acuminata var. montana (Arecaceae), and subjected them to barrier-protected and wind-exposed treatments. For C. erosum , we compared the allometry of stems and branches against three allometric models that predict that plant height or branch length increases at the 1, 2/3, and 1/2 power of stem diameter. Only the geometric similarity model (scaling exponent of 1) seemed to hold when plants were exposed to the wind. We found relatively fewer leaves per number of branches produced and fewer leaves per increment of branch diameter in the plants of C. erosum exposed to the wind. Mean petiole length ratios (petiole length/basal radius) of P. acuminata were higher on leaves of barrier-protected plants for both simple and compound leaves, indicating that petioles were stouter and mechanically safer in the wind-exposed plants. We suggest that alteration of the allometric relationships of plant parts, organs, or plant modules (stems and branches of C. erosum and leaves of P. acuminata ) and alteration of the number of plant parts (leaves and branches of C. erosum ) are adaptive responses resulting from the mechanical perturbation induced by wind in the elfin forest.     

祁连山北坡霸王枝-叶性状关系的个体大小差异

枝与叶片权衡关系的个体大小差异性, 是物种形成不同冠层结构充分利用空间资源的一种策略, 有利于植株通过构型调整自身的光合效率和增强竞争力。在祁连山北坡荒漠草地, 根据体积将霸王(Zygophyllum xanthoxylum)分为3个大小等级(I级: (植株体积的立方根(d) ≤ 60 cm)、II级(60 cm < d ≤ 120 cm)和III级(d > 120 cm)), 采用标准化主轴估计方法, 研究了不同大小等级霸王种群枝长度与叶面积、叶数量的生长关系。结果表明: 随着植株大小等级增大, 霸王的枝长度、叶面积、枝横截面积逐渐增大, 叶数量呈逐渐减小趋势; 霸王枝长度的增长速度大于或等于叶面积的增长速度, 枝长度的增加速度大于叶数量的增加速度; 随着个体大小等级的增加, 霸王枝长度与叶面积的异速斜率、枝长度与叶数量回归方程的y轴截距均显著减小, 即植株叶面积与枝长度比值、叶数量的投入均显著降低。为提高资源利用效率, 霸王小个体植株倾向于短枝上着生大量的小叶, 大个体植株趋向于长枝上着生少量的大叶, 随植株大小等级增加, 叶面积增加而叶数量降低, 对小枝的资源配置具有一定的影响。     

Influence of host plant heterogeneity on the distribution of a birch aphid

Abstract.  1. The spatial and temporal abundance of the aphid Euceraphis betulae was investigated in relation to heterogeneity in host plant ( Betula pendula ) vigour and pathogenic stress. The performance of aphids feeding on vigorous and stressed foliage was also examined.
2. The plant stress and plant vigour hypotheses have been suggested as opposing ways in which foliage quality influences herbivore abundance. In many plants, however, vigorous growing foliage co-exists with stressed or damaged foliage.
3. There was a negative correlation between branch growth (vigour) and branch stress (leaf chlorosis), with the most vigorous branches displaying little or no stress, and the most stressed branches achieving poor growth. There was a similar negative correlation between vigour and stress at the level of individual trees, which themselves represented a continuum in quality.
4. At the beginning of the season, E. betulae were intermittently more abundant on vigorous branches than on branches destined to become stressed, but aphids became significantly more abundant on stressed branches later in the season, when symptoms of stress became apparent. Similar patterns of aphid abundance were seen on vigorous and stressed trees in the following year.
5. Euceraphis betulae performance was generally enhanced when feeding on naturally stressed B. pendula leaves, but there was some evidence for elevated potential reproduction when feeding on vigorous leaves too.
6. Overall, plant stress probably influences E. betulae distribution more than plant vigour, but the temporal and spatial variability in plant quality suggests that plant vigour could play a role in aphid distribution early in the season.     

Interactive effects of elevated C02 and temperature on the anatomical characteristics of leaves in eleven species

The anatomical features of leaves in 11 species of plants grown in a temperature gradient and a temperature + CO2 gradient were studied.The palisade parenchyma thickness,the spongy parenchyma thickness and the total leaf thickness were measured and analyzed to investigate the effects of elevated temperature and CO2 on the anatomical characteristics of the leaves.Our results show that with the increase of temperature,the leaf thickness of C4 species increased while the leaf thickness of C3 species showed no constant changes.With increased CO2,seven out of nine C3 species exhibited increased total leaf thickness.In C4 species,leaf thickness decreased.As for the trend on the multi-grades,the plants exhibited linear or non-linear changes.With the increase of temperature or both temperature and CO2 for the 11 species investigated,leaf thickness varied greatly in different plants (species) and even in different branches on the same plant.These results demonstrated that the effect of increasing CO2 and temperature on the anatomical features of the leaves were species-specific.Since plant structures are correlated with plant functions,the changes in leaf anatomical characteristics in elevated temperature and CO2 may lead to functional differences.     

Ontogenetic approach to the assessment of plant resistance to prolonged stress using chlorophyll fluorescence induction method

The review deals with objective reasons that limit the use of chlorophyll fluorescence induction kinetics (Chl FIK) method in plant ecology. Based on the ontogenetic approach (analysis and comparison of the dynamics of the Chl fluorescence ratio F_p/F_s and physiological characteristics of plant leaves in ontogeny) possible criteria for the estimation of general plant resistance of photosynthetic apparatus (PSA) to prolonged stress affects are proposed. One of these criteria is the presence or absence of a steady-state phase in the dynamics of Chl fluorescence ratio F_p/F_s (or R_fd) of plant leaves after they stop growing. We also propose to use the duration of the steady-state phase and variability of Chl fluorescence ratios R_fd and F_p/F_s in this period for quantitative assessment of plant PSA resistance to prolonged stress during plant leaf ontogeny. The paper is dedicated to memory of Prof. F.Ya. Sid’ko.     

散孔材与环孔材树种枝干、叶水力学特性的比较研究

为揭示散孔材与环孔材树种树木水分生理特性的差异,选取了常见的3种散孔材落叶树种(毛白杨、法国梧桐和樱花)和3种环孔材落叶树种(刺槐、合欢和白蜡),研究了其枝干与叶水力学性质的差异及其协调性。结果表明:3种环孔材树种枝干横截面积基础上的最大比导水率(Ks-max)大于3种散孔材树种,但其木质部对空穴化的脆弱性(P50branch)高于散孔材树种,6种树木枝干的水分传输能力和抵抗空穴化能力之间存在一种相互制约的权衡关系。3种散孔材与3种环孔材树种的叶最大水力导度(Kl-max)和水力脆弱性(P50leaf)并无显著差异;对于3种散孔材树种,叶的水力脆弱性要高于枝干,但对3种环孔材树种而言,枝干的水力脆弱性要高于叶。6种树木枝干和叶的水力学性质(Kmax、P50)之间并无相关关系。这些结果表明:散孔材与环孔材树种的枝干水力学特性有明显差异,但叶水力学特性无差异;枝干与叶水力学性质之间是相互独立的。     

Branch development controls leaf area dynamics in grapevine (Vitis vinifera) growing in drying soil

BACKGROUND AND AIMS: Soil water deficit is a major abiotic stress with severe consequences for the development, productivity and quality of crops. However, it is considered a positive factor in grapevine management (Vitis vinifera), as it has been shown to increase grape quality. The effects of soil water deficit on organogenesis, morphogenesis and gas exchange in the shoot were investigated. METHODS: Shoot organogenesis was analysed by distinguishing between the various steps in the development of the main axis and branches. Several experiments were carried out in pots, placed in a greenhouse or outside, in southern France. Soil water deficits of various intensities were imposed during vegetative development of the shoots of two cultivars ('Syrah' and 'Grenache N'). KEY RESULTS: All developmental processes were inhibited by soil water deficit, in an intensity-dependent manner, and sensitivity to water stress was process-dependent. Quantitative relationships with soil water were established for all processes. No difference was observed between the two cultivars for any criterion. The number of leaves on branches was particularly sensitive to soil water deficit, which rapidly and strongly reduced the rate of leaf appearance on developing branches. This response was not related to carbon availability, photosynthetic activity or the soluble sugar content of young expanding leaves. The potential number of branches was not a limiting factor for shoot development. CONCLUSIONS: The particularly high sensitivity to soil water deficit of leaf appearance on branches indicates that this process is a major determinant of the adaptation of plant leaf area to soil water deficit. The origin of this particular developmental response to soil water deficit is unclear, but it seems to be related to constitutive characteristics of branches rather than to competition for assimilates between axes differing in sink strength.     

How do Plant Morphological Characteristics,Species Composition and Richness Regulate Eco-hydrological Function?

Although considerable research has focused on the relationship between ecosystem structure and function, interactions of plant morphological characteristics, species composition and richness with eco-hydrological functions remain unclear. We measured water adherence (i.e. the capacity of a plant species to retain water), documented plant surface morphology and observed surface runoff at three sites in China. The adhering water ratios for each plant species differed, ranging from 17.1% to 151.5% in leaves, and from 14.4% to 41.1% in branches. Small, light-weight, soft, non-cuticularized leaves that were densely situated on small branches showed good water adherence. The next best adherence was found by branches with intermediately coarse surfaces. The plant species with high standing biomass also showed good water adherence, and the contribution of a species to total adherence was dependent upon its aboveground standing biomass. Vegetation parameters strongly affected water adherence, whereas the effect of species richness was not significant. Conversely, species richness showed a significant influence on surface runoff. The effect of plant morphological characteristics and composition constitutes a basic process in the regulation of eco-hydrological function, and vegetation parameters play somewhat different roles in that regulation. The key roles must therefore be considered from a management perspective.