Feild T S, Lee D W, Holbrook N M
Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA 02138, USA.
Plant Physiol. 2001 Oct;127(2):566-74.
Why the leaves of many woody species accumulate anthocyanins prior to being shed has long puzzled biologists because it is unclear what effects anthocyanins may have on leaf function. Here, we provide evidence for red-osier dogwood (Cornus stolonifera) that anthocyanins form a pigment layer in the palisade mesophyll layer that decreases light capture by chloroplasts. Measurements of leaf absorbance demonstrated that red-senescing leaves absorbed more light of blue-green to orange wavelengths (495-644 nm) compared with yellow-senescing leaves. Using chlorophyll a fluorescence measurements, we observed that maximum photosystem II (PSII) photon yield of red-senescing leaves recovered from a high-light stress treatment, whereas yellow-senescing leaves failed to recover after 6 h of dark adaptation, which suggests photo-oxidative damage. Because no differences were observed in light response curves of effective PSII photon yield for red- and yellow-senescing leaves, differences between red- and yellow-senescing cannot be explained by differences in the capacities for photochemical and non-photochemical light energy dissipation. A role of anthocyanins as screening pigments was explored further by measuring the responses PSII photon yield to blue light, which is preferentially absorbed by anthocyanins, versus red light, which is poorly absorbed. We found that dark-adapted PSII photon yield of red-senescing leaves recovered rapidly following illumination with blue light. However, red light induced a similar, prolonged decrease in PSII photon yield in both red- and yellow-senescing leaves. We suggest that optical masking of chlorophyll by anthocyanins reduces risk of photo-oxidative damage to leaf cells as they senesce, which otherwise may lower the efficiency of nutrient retrieval from senescing autumn leaves.
为何许多木本植物的叶片在脱落之前会积累花青素,这一直困扰着生物学家,因为尚不清楚花青素可能对叶片功能产生何种影响。在此,我们为红枝山茱萸(Cornus stolonifera)提供了证据,表明花青素在栅栏叶肉层形成色素层,减少了叶绿体对光的捕获。叶片吸光度测量表明,与变黄的叶片相比,变红衰老的叶片吸收更多蓝绿色至橙色波长(495 - 644纳米)的光。通过叶绿素a荧光测量,我们观察到,高光胁迫处理后,变红衰老叶片的最大光系统II(PSII)光子产量恢复了,而变黄的叶片在暗适应6小时后未能恢复,这表明存在光氧化损伤。由于在变红和变黄衰老叶片的有效PSII光子产量的光响应曲线中未观察到差异,变红和变黄衰老叶片之间的差异无法用光化学和非光化学光能耗散能力的差异来解释。通过测量PSII光子产量对蓝光(优先被花青素吸收)与红光(吸收较差)的响应,进一步探究了花青素作为筛选色素的作用。我们发现,暗适应的变红衰老叶片在蓝光照射后PSII光子产量迅速恢复。然而,红光在变红和变黄衰老叶片中均诱导了类似的、PSII光子产量的长时间下降。我们认为,花青素对叶绿素的光学遮蔽降低了叶片细胞衰老时光氧化损伤的风险,否则可能会降低从衰老秋叶中回收养分的效率。