异质性光对野牛草叶片解剖结构的影响

doi:10.17521/cjpe.2015.0377

摘要/Abstract

摘要：

自然界中植物生长所需资源通常呈异质性分布, 具有发达匍匐茎的野牛草(Buchloe dactyloides)在蔓延过程中相连克隆分株常生活在异质性的光环境中。有研究证明, 在异质性光条件下, 植株幼叶的叶片解剖结构受成熟叶所处光照条件的影响, 而异质性光条件下克隆分株的叶片形态解剖结构是否也受相连分株所处光照条件的影响则未见报道。通过设置高光(全光照)和低光(遮阴)两个水平, 对由匍匐茎相连的野牛草克隆分株施以同质和异质性光处理, 研究了异质性光对野牛草叶片解剖结构的影响。结果发现: 在异质性光环境中, 遮阴的野牛草克隆分株的主脉直径、维管束鞘细胞个数、叶片厚度以及近轴侧和远轴侧叶肉细胞的厚度均显著降低; 同质性的低光处理对这些指标则没有显著影响。在异质性光处理下, 未遮阴姊株近轴侧和远轴侧叶肉细胞的厚度以及远轴侧的气孔大小显著增加, 而未遮阴的妹株近轴侧和远轴侧叶肉细胞的厚度、气孔密度和气孔大小、叶片厚度和维管束鞘细胞个数则会降低。同质高光处理下克隆分株近轴侧和远轴侧的气孔密度和气孔大小显著高于同质低光。野牛草克隆分株近轴侧和远轴侧叶肉细胞的厚度、气孔密度和气孔大小受相连分株所处光照条件的显著影响。该研究结果表明: 未遮阴的姊株因为与遮阴的妹株相连而显著受益, 而未遮阴的妹株则因为与遮阴的姊株相连而损耗严重; 在异质性光处理下, 遮阴分株叶片形态上缩减的可塑性生长是为减少维持其存活的消耗, 提高遮阴分株存活率的一种适应性表现。

关键词: 野牛草, 异质性光, 解剖结构, 气孔密度

Abstract: Aims

Essential resources for plant growth are usually patchily distributed. During the process of propagation, interconnected ramet pairs of stoloniferous plant buffalograss (Buchloe dactyloides) may therefore experience contrasting resource supply such as light. Under heterogeneous light supply, anatomical structure of newly developed leaves is regulated by the light condition of mature leaves. However, little is known about whether leaf anatomical structure of clonal ramets is affected by the light environment of interconnected ramets in clonal plants under heterogeneous light supply.

Methods

Two light levels were set, with high light (natural sunlight) and low light (shade, 10% natural sunlight). Interconnected ramet pairs of buffalograss were exposed to homogeneous or heterogeneous light conditions.

Important findings Main vein diameter, bundle sheath cell number, leaf thickness and adaxial/abaxial mesophyll thickness of shaded ramets were remarkably decreased in spatially heterogeneous light environment; while no signifiacnt difference of these parameters was observed between ramets developed under homogeneous high light and low light conditions. Under heterogeneous light supply, adaxial/abaxial mesophyll thickness and abaxial stomatal size in unshaded elder daughter ramets (EDR) were remarkably increased, while adaxial/abaxial mesophyll thickness, stomatal density and size, leaf thickness, and bundle sheath cell number in unshaded younger daughter ramets (YDR) were reduced. Ramets under homogeneous high light conditions had higher stomatal density and larger stomata than those under homogeneous low light conditions. Adaxial/abaxial mesophyll thickness, stomatal density and size in buffalograss ramets were significantly affected by the light conditions of interconnected ramets.

Conclusions

Unshaded EDR benefit from their connection to shaded YDR, while unshaded YDR experience marked cost due to its connection to shaded EDR. The plastic decrease of shaded ramets under heterogeneous light may be associated with its reduced survival cost, and elevated survival rate under shading.

Key words: buffalograss, light heterogeneity, anatomical structure, stomatal density

郝晨淞, 王庆凯, 孙小玲. 异质性光对野牛草叶片解剖结构的影响. 植物生态学报, 2016, 40(3): 246-254. DOI: 10.17521/cjpe.2015.0377

Chen-Song HAO, Qing-Kai WANG, Xiao-Ling SUN. Effects of light heterogeneity on leaf anatomical structure in Buchloe dactyloides. Chinese Journal of Plant Ecology, 2016, 40(3): 246-254. DOI: 10.17521/cjpe.2015.0377

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2015.0377

https://www.plant-ecology.com/CN/Y2016/V40/I3/246

图/表 7

图1 试验开始时由匍匐茎连接的野牛草母株、姊株和妹株。

Fig. 1 Diagrammatic presentation of a Buchloe dactyloides stolon segment consisting of mother ramet, elder daughter ramet and younger daughter ramet at the start of this experiment.

表1 姊妹分株接受的不同光处理

Table 1 The pattern of light treatments to elder daughter ramets (EDR) and younger daughter ramets (YDR)

试验处理 Treatment	姊株 EDR	妹株 YDR
HH	高光 High light	高光 High light
HL	高光 High light	低光 Low light
LH	低光 Low light	高光 High light
LL	低光 Low light	低光 Low light

图2 光学显微镜下野牛草叶片的横切结构图。近轴侧和远轴侧叶肉细胞厚度的测定参考Jiang等(2011)的方法, 选取维管束中央(叶片中间)的位置拉线, 如上图的虚线所示。a, 叶肉细胞; b, 维管束鞘细胞; c, 厚壁组织; d, 表皮细胞; e, 木质部; f, 韧皮部; g, 泡状细胞。

Fig. 2 Illustration of an image used to measure the adaxial (lower) and abaxial (upper) mesophyll tissues of Buchloe dactyloides in a cross-section light micrograph. The adaxial and abaxial mesophyll thickness were measured separately relative to the middle of the bundle sheath as shown by the dashed line, which generally represented the middle of the leaf (Jiang et al., 2011). a, mesophyll cells; b, bundle sheath cells; c, sclerenchyma; d, epidermal cell; e, xylem; f, phloem; g, motor cell.

图3 同质和异质性光处理下野牛草姊株和妹株的主脉直径(A)、维管束鞘细胞个数(B)和维管束鞘细胞与叶肉接触面的长度(C) (平均值±标准误差, n = 7)。不同小写/大写字母表示处理间差异显著(p < 0.05)。HH、HL、LH、LL同表1。

Fig. 3 Main vein diameter (MVD) (A), bundle sheath cell number (BSCN) (B), and total contact length between bundle sheath and mesophyll cells (CLBM) (C) of Buchloe dactyloides elder daughter ramets (EDR) and younger daughter ramets (YDR) under homogeneous and heterogeneous light treatments (means ± SE, n = 7). Different letters indicate significant difference among treatments (p < 0.05). For HH, HL, LH and LL, see Table 1.

图4 同质和异质性光处理下野牛草姊株和妹株的叶片厚度(A)、近轴侧叶肉细胞的厚度(B)和远轴侧叶肉细胞的厚度(C) (平均值±标准误差, n = 7)。不同小写/大写字母表示处理间差异显著(p < 0.05)。HH、HL、LH、LL同表1。

Fig 4 Leaf thickness (LT) (A), adaxial mesophyll thickness (AdMT) (B), and abaxial mesophyll thickness (AbMT) (C) of Buchloe dactyloides elder daughter ramets (EDR) and younger daughter ramets (YDR) under homogeneous and heterogeneous light treatments (means ± SE, n = 7). Different letters indicate significant difference among treatments (p < 0.05). For HH, HL, LH and LL, see Table 1.

表2 年龄、分株自身所处光照条件和相连分株所处光照条件及其交互作用对野牛草叶片解剖特征的三因素方差分析

Table 2 F-values of three-way ANOVA for the effects of age (A), local condition (Lc), remote condition (Rc), and their interactions on leaf anatomical traits of Buchloe dactyloides

处理 Treatment	自由度 Degree of freedom	主脉直径 MVD	维管束鞘细胞个数 BSCN	维管束鞘细胞与叶肉接触面的长度 CLBM	叶片厚度 LT	近轴侧叶肉细胞厚度 AdMT	远轴侧叶肉细胞厚度 AbMT	近轴侧气孔密度 AdSD	远轴侧气孔密度 AbSD	近轴侧气孔大小 AdSZ	远轴侧气孔大小 AbSZ
年龄 A	1, 48	0.754	0.388	0.017	1.168	4.19^*	4.28^*	0.368	0.116	33.925^***	22.998^***
分株自身所处光照 Lc	1, 48	10.667^**	0.028	1.864	4.027	0.910	3.740	23.540^***	31.030^***	55.845^***	85.684^***
相连分株所处光照 Rc	1, 48	3.265	0.447	0.209	0.559	8.380^**	17.550^***	29.793^***	35.030^***	130.628^***	98.520^***
A × Lc	1, 48	0.018	2.495	0.988	0.008	0.110	1.630	0.828	0.485	37.874^***	102.998^***
Lc × Rc	1, 48	11.227^**	28.096^***	6.339^*	10.221^**	16.570^***	20.700^***	4.506^*	1.486	2.041	0.112
A × Rc	1, 48	0.883	4.457^*	0.001	1.380	12.780^***	3.730	0.092	0.059	21.304^***	81.604^***
A × Lc × Rc	1, 48	0.285	2.592	1.299	0.079	1.390	3.490	0.368	0.021	0.592	66.281^***

图5 同质和异质性光处理下野牛草姊株和妹株近轴侧和远轴侧的气孔密度(A和B)以及近轴侧和远轴侧的气孔大小(C和D) (平均值±标准误差, n = 7)。不同小写/大写字母表示处理间差异显著(p < 0.05)。HH、HL、LH、LL同表1。

Fig. 5 Adaxial stomatal density (AdSD) (A), abaxial stomatal density (AbSD) (B), adaxial stomatal size (AdSZ) (C) and abaxial stomatal size (AbSZ) (D) of Buchloe dactyloides elder daughter ramets (EDR) and younger daughter ramets (YDR) under homogeneous and heterogeneous light treatments (means ± SE, n = 7). Different letters indicate significant difference among treatments (p < 0.05). For HH, HL, LH and LL, see Table 1.

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