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植物叶片秋季衰老会导致叶色变化,暖温带落叶林最显著的特征是秋季叶色丰富[1-2]。目前植物叶片呈色机制仍不完全清楚[3]。研究发现叶色与叶片色素种类、含量和分布有关[4-5],叶片呈色的直接原因是叶片中色素种类和比例的变化[6]。大多数植物叶片叶绿素含量秋季开始时缓慢下降,然后迅速下降,同时叶片组织中开始积累花青素[7-9],秋季叶色变红主要与叶片中花青素积累有关[7,10-11]。研究表明叶片色素含量与色差参数间具有一定相关性,如花色素苷和叶绿素含量的多少决定着a*值大小[12]。叶片花色苷的主要功能是使叶片免受光合抑制影响[13],同时有利于叶片营养元素的吸收再利用[14]。与N含量较高的叶片相比,N含量较低的叶片更早且更完全变红[4],表明色素与营养元素在叶片中的变化高度相关[15]。N和P是植物生长发育的限制性元素[16],同时也是各种蛋白质和遗传物质的重要组成元素[17]。因此,利用色差参数对主要色素含量进行预测,了解营养元素动态和叶色在叶片衰老期的变化关系,有助于了解植物变色期的适应策略,进一步揭示植物叶片秋冬季呈色规律[18]。
北美栎树(Quercus spp.)引种至我国已有20多年,其中有些树种引种点较多,种植广泛,生长良好[19]。作为观赏树种,北美栎树特别是红栎组(Sect. Lobatae)栎树秋冬季叶色丰富。研究表明平均气温是引起纳塔栎(Q. texana Buckley)秋季叶片变色的主要因素,其次是最高气温[20]。姜琳等探讨了纳塔栎、舒玛栎(Q. shumardii Buckland)等树种在秋冬转色期叶色变化的生理机制[21],但较少涉及关于秋季色素含量,特别是营养元素与叶色量化参数间变化关系方面的研究。由于栎树资源丰富,不同树种之间秋季叶色差异显著,这一现象可能与其秋季花青素积累差异显著有关[11,22]。同时,不同栎树叶片营养元素变化的差异可能也间接导致其叶色显著不同。因此,本研究选择从北美引种的7种栎树以及乡土树种白栎(Q. fabri Hance)为研究对象,通过研究8种栎树叶片变色期有关生理指标变化,进而揭示栎树叶片营养元素生态化学计量特征、色素和叶色参数间的关系,以期为栎树园林绿化应用和彩叶新品系选育提供科学依据。
栎树秋季叶色多样性及相关生理特性的变化
Leaf Color Diversity and the Change of Physiological Characteristics of Quercus spp. During the Discoloration Period in Autumn
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摘要:
目的 研究栎树叶片变色期相关生理物质含量的变化,探索其叶色多样性与生理特征变化间的内在联系。 方法 以红栎组的纳塔栎、舒玛栎、柳叶栎、月桂叶栎、樱皮栎,白栎组的琴叶栎、牛栎以及白栎等8种栎属植物4 年生苗为试材,对其色差参数(L*、a*、b*)、色素、可溶性糖和营养元素含量进行测定分析。 结果 秋季栎树叶片叶色参数L*值变化幅度小,较为稳定。色差参数a* 值、b*值、叶绿素含量和类胡萝卜素含量呈下降趋势,花青素含量则逐渐上升;N、P含量在叶片变色期逐渐降低,而可溶性糖含量增加。同时a*值与花青素、可溶性糖含量、C∶N和C∶P呈极显著正相关,与叶绿素、类胡萝卜素、N和P含量呈极显著负相关。通常红栎组栎树叶色较白栎组红。由于各树种各指标变化幅度不同,导致不同树种叶色差异显著,且叶色保持期不同。叶片变色后期各栎树叶色从绿色变为黄红色、深红色、鲜红色或棕色。其中纳塔栎秋季叶色最红,同时呈现红色的时间最长。 结论 红栎组栎树较早进入变色期;变色前期各指标变化幅度小,此后叶绿素和类胡萝卜素被快速分解,花青素大量合成,是影响栎树秋季叶色的关键因素;变色期叶片N和P含量逐渐下降且利用率显著提高。 Abstract:Objective To study the change of physiological characteristics changes of Quercus spp. during the discoloration period and found the relationship between leaf color diversity and physiological changes. Method 4-year-old seedlings of Sect. Lobatae (Quercus texana, Q. shumardii, Q. phellos, Q. laurifolia and Q. pagoda) and Sect. Quercus (Q. lyrata, Q. michauxii and Q. fabri) were used to measure chromatic aberration parameters (L*, a*, and b*), pigment, soluble sugar, C, N and P contents during the leaf senescence stage. Result During the autumn, the leaf senescence period, the variation range of L* value was small and stable. The values of a* and b*, chlorophyll and carotenoid contents decreased whereas the anthocyanin content increased gradually. The leaf N and P contents decreased gradually in the senescing leaves, but the contents of soluble sugar in leaves increased. The a* value showed significantly positive correlations with anthocyanin and soluble sugar content, C:N ratio and C:P ratio; and negative correlations with chlorophyll, carotenoid, N and P content. In general, the leaf color of Sect. Lobatae was redder than that of Sect. Quercus, while there were significant differences among species in terms of leaf color and leaf color retention period. Due to the different magnitude of variation in each index of each species, the leaf color of Quercus spp. changed from green to yellow-red, crimson, bright red or brown in the late stage of leaf discoloration. The leaf color of Q. texana was the reddest in autumn and the red leaves occurred the longest. Conclusion Usually, the leaf color of Sect. Lobatae will change earlier than that of Sect. Quercus. In the early stages, the variation of parameters is very small. After, the leaf color of Quercus spp. is mostly affected by chlorophyll and carotenoids break down and anthocyanins will synthesize quickly. The N and P contents will gradually decrease, and the utilization rate increase significantly. -
Key words:
- Quercus spp.
- / leaf color
- / anthocyanin
- / nutrients
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