竹笋期竹箨和笋体的日间蒸腾特性及其对水分运输的影响

doi:10.17521/cjpe.2021.0164

植物生态学报 ›› 2021, Vol. 45 ›› Issue (12): 1365-1379.DOI: 10.17521/cjpe.2021.0164

竹笋期竹箨和笋体的日间蒸腾特性及其对水分运输的影响

李唐吉¹, 王懋林¹^,², 曹颖¹^,^*(), 徐刚¹, 杨琪祺¹, 任思源¹, 胡尚连¹^,²^,^*()

¹西南科技大学生命科学与工程学院, 四川绵阳 621010
²四川省生物质资源利用与改性工程技术研究中心, 四川绵阳 621010

收稿日期:2021-04-28 接受日期:2021-07-15 出版日期:2021-12-20 发布日期:2021-09-18
通讯作者: 曹颖,胡尚连
作者简介:Hu SL, hushanglian@126.com
^*Cao Y, caoying@swust.edu.cn;
基金资助:
四川省“十四五”重点攻关项目(20zs2172);四川省重点研发项目(2019YFN0005);四川省大学生创新创业训练计划资助项目(19xcy056)

Diurnal transpiration of bamboo culm and sheath and their potential effects on water transport during the bamboo shoot stage

LI Tang-Ji¹, WANG Mao-Lin¹^,², CAO Ying¹^,^*(), XU Gang¹, YANG Qi-Qi¹, REN Si-Yuan¹, HU Shang-Lian¹^,²^,^*()

¹School of Life Science and Engineering, Southwest University of Science and Technology, Mianyang, Sichuan 621010, China
²Engineering Research Center for Biomass Resource Utilization and Modification of Sichuan Province, Mianyang, Sichuan 621010, China

Received:2021-04-28 Accepted:2021-07-15 Online:2021-12-20 Published:2021-09-18
Contact: CAO Ying,HU Shang-Lian
Supported by:
14th Five-Year Key Project of Sichuan Province(20zs2172);Key Research and Development Projects of Sichuan Province(2019YFN0005);Sichuan’s Training Program of Innovation and Enterpreneurship for Undergraduate(19xcy056)

摘要/Abstract

摘要：

竹子的高速生长主要发生在无枝无叶的笋期, 并对水分需求巨大。水分不仅参与植物体内各种代谢, 而且水分转运可促进光合产物、矿质元素、生长激素等物质流动。竹子夜间主要由根压驱动水分转运, 但在日间尤其是下午根压基本为负值, 明确竹笋日间蒸腾作用发生机制及其对水分运输的影响对竹林培育有重要意义。该研究以不同伸长阶段的慈竹(Bambusa emeiensis)笋为材料, 研究了茎秆和竹箨的气孔特征、气孔导度与蒸腾速率等生理特征及在离体条件下竹笋的水分转运速率。结果表明: 1)不同发育状态的竹笋茎秆及箨鞘表面均分布有大量气孔, 气孔小且凹陷, 光合速率及叶绿素a、b含量极低, 但气孔导度和蒸腾速率均显著高于成熟叶片, 表明笋体和箨鞘是竹笋主要的呼吸和蒸腾部位。2)离体条件下竹笋的番红示踪表明, 高生长阶段的竹笋茎秆中番红上升速率较快, 有着较强的蒸腾。竹箨分离后, 番红仍然能够扩散和运输, 表明笋体茎秆也存在一定的蒸腾, 但与竹箨包裹的竹笋相比, 番红在分离竹箨后的笋体中上升速度显著下降, 表明竹箨对笋体内水分运输影响较大。3)箨环处的组织解剖发现, 节间的纵向维管束在竹节处特化形成一个类板状结构, 弯曲伸入竹箨, 是竹箨影响笋体内水分运输的重要结构基础。上述结果表明, 日间竹笋水分通过茎秆和竹箨表面的气孔大量蒸散, 产生蒸腾拉力驱动笋体内水分转运。该研究也发现, 随着茎秆成熟, 竹箨松动并开始脱落, 茎秆表面的气孔宽度增加, 加大了气孔的开口大小, 增大了节间气孔与大气水气交换的有效面积, 在一定程度上弥补了竹箨脱落时减少的蒸腾拉力。

关键词: 日间蒸腾, 水分运输, 气孔, 竹箨, 竹笋

Abstract:

Aims Water transport is of critical impact on the growth of bamboo shoots. The mechanism of diurnal water transpiration by bamboo shoots, and its effect on water transport was not yet fully comprehended. The objectives of this study include: 1) to characterize the structure of bamboo shoots that influence transpiration; 2) to examine the ability of the structure in transporting water; 3) to examine the structure of culm sheath for understanding water transfer in bamboo shoots.

Methods Bambusa emeiensisshoots of different elongation stages were used to characterize the stomata, for examining stomatal conductance (G_s), transpiration rate (T_r) and other physiological features of culm sheaths and shoot body. Water transport rate of shoots in vitro was also examined.

Important findings The results showed that: 1) there are a large number of small and concave stomata on the surface of culm sheath and bamboo shoot of different development states, their net photosynthetic rate (P_n), chlorophyll (Chl) a, and Chl b content were extremely low, but G_s and T_r were significantly higher than that of the mature leaves. These indicated that culm sheath and bamboo shoot play important roles in transpiration and respiration. 2) The tracing results of safranine solution in vitro showed that shoots of higher daily elongation rate had stronger transpiration, and were higher rising rate of safranine solution in the bamboo shoots. When culm sheaths of bamboo shoots were separated, the safranine solution could still diffused and transported in the shoot bodies, indicating that the shoot body itself carried some transpiration. However, compared with the intact bamboo shoots wrapped by culm sheaths, the rising safranine solution decreased significantly after the culm sheaths were separated, indicating that the culm sheath could make important contribution for the water transportation. 3) Anatomical analysis showed that some longitudinal vascular bundles of the internodes formed a plate-like structure (PLS) at the bamboo nodes, and the ends of PLS gradually bent and stretched into the bamboo sheath. The PLS may be an important structural basis for water transport of bamboo sheath.

Key words: diurnal transpiration, water transport, stomata, culm sheath, bamboo shoot

李唐吉, 王懋林, 曹颖, 徐刚, 杨琪祺, 任思源, 胡尚连. 竹笋期竹箨和笋体的日间蒸腾特性及其对水分运输的影响. 植物生态学报, 2021, 45(12): 1365-1379. DOI: 10.17521/cjpe.2021.0164

LI Tang-Ji, WANG Mao-Lin, CAO Ying, XU Gang, YANG Qi-Qi, REN Si-Yuan, HU Shang-Lian. Diurnal transpiration of bamboo culm and sheath and their potential effects on water transport during the bamboo shoot stage. Chinese Journal of Plant Ecology, 2021, 45(12): 1365-1379. DOI: 10.17521/cjpe.2021.0164

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0164

https://www.plant-ecology.com/CN/Y2021/V45/I12/1365

图/表 9

图1 慈竹不同发育阶段的箨鞘、箨叶及成熟叶片的气孔形态。A, D, 幼嫩和成熟竹箨, 虚线以上为箨叶, 虚线以下为箨鞘。B, E, 扫描电镜下幼嫩及成熟箨鞘的气孔, 右上图为气孔放大图。C, F, 扫描电镜下幼嫩及成熟箨叶的气孔。G-I, 竹子成熟叶片的形态(G)和气孔特征(H, I)。箭头指向气孔。

Fig. 1 Stoma morphology of culm sheath, sheath leaf and mature branch leaf at different developmental stages of Bambusa emeiensis. A, D, Culm sheaths at the young and mature stage, respectively, the culm sheaths and sheath leaf were represented at above and below the dotted line, respectively. B, E, The stomata of the young and mature culm sheath respectively under a scanning electron microscope. The figures at upper right corner of B and E show the enlarged stomata. C, F, The stomata of young and mature sheath leaves respectively under scanning electron microscope. G-I, Leaf morphology (G) and stoma (H, I) of mature branch leaf. Arrows point to stoma.

表1 慈竹不同发育阶段箨鞘、箨叶及成熟叶片的气孔大小和密度(平均值±标准差)

Table 1 Stoma size and density of culm sheath, sheath leaf and mature branch leaf at different stages of Bambusa emeiensis (mean ± SD)

	箨鞘 Culm sheath		箨叶 Sheath leaf		成熟叶片 Mature leaf
	幼嫩 Young	成熟 Mature	幼嫩 Young	成熟 Mature	成熟叶片 Mature leaf
气孔长度 Stomatal length (μm)	5.73 ± 0.37^d	7.83 ± 0.42^c	10.53 ± 0.59^b	12.14 ± 0.34^a	12.07 ± 0.63^a
气孔宽度 Stomatal width (μm)	2.07 ± 0.16^b	2.30 ± 0.20^a	0.96 ± 0.07^c	1.13 ± 0.05^c	1.12 ± 0.05^c
气孔密度 Stomatal density (mm^-2)	425.40 ± 32.41^b	346.72 ± 14.57^c	593.31 ± 28.75^a	589.70 ± 31.01^a	561.70 ± 34.20^a
气孔长/宽 Stomatal length-to-width	2.85 ± 0.22^c	3.40 ± 0.21^b	10.41 ± 0.87^a	10.60 ± 0.65^a	10.71 ± 0.77^a

图2 慈竹不同发育阶段笋体(茎秆)的气孔形态。A-F, 不同高度竹笋茎秆基部形态。基部第一节间正在伸长(A, B); 基部第一节间完全伸长, 节间部分露出竹箨(C, D), 箭头指向节间露出位置(C); 基部第一节间成熟, 竹箨松动并开始脱落(E, F), 箭头指向竹箨松动处(E)。B, D, F中的方框为取样部位, 箭头指向气孔, 虚线以上为无根节, 虚线以下为有根节。G-L, 扫描电镜下竹笋茎秆基部第一节间的气孔形态。G-I分别为节间的伸长期, 完全伸长期和成熟期; J-L分别为G-I中方框位置的局部放大图。

Fig. 2 Stoma morphology of shoot body (culm) at different development stages of Bambusa emeiensis. A-F, the morphology of basal bamboo shoots with different height, in elongating (A, B), elongated fully (C, D), and mature (E, F) stages respectively in the first basal internodes. Above the dotted line are rootless nodes, below the dotted line are rooted nodes (B, D, F). The arrow in C points to the culm exposed from the culm sheath; arrow in E, to the loose position of the culm sheath. G-L, the stomatal morphology, under scanning electron microscope, in elongating (G), elongated fully (H), and mature (I) stages respectively in the first basal internode, the arrow points to the stomata. J-L, enlarged pictures of the boxes in G-I, respectively.

表2 慈竹不同发育阶段笋体(茎秆)气孔大小和密度(平均值±标准差)

Table 2 Size and density of stoma in shoot body (culm) at different developmental stages of Bambusa emeiensis (mean ± SD)

基部第一节间 Basal first internode	伸长期 Elongating	完全伸长期 Fully elongated	成熟期 Mature
节间长 Internode length (cm)	4.50 ± 0.20^b	12.43 ± 0.55^a	12.40 ± 0.92^a
节间宽 Internode width (cm)	3.56 ± 0.35^b	7.46 ± 0.15^a	7.43 ± 0.25^a
节间表面 Internode surface area (cm²)	48.44 ± 6.12^b	279.69 ± 9.53^a	278.01 ± 25.34^a
气孔长度 Stoma length (μm)	1.60 ± 0.15^b	3.43 ± 0.31^a	3.46 ± 0.21^a
气孔宽度 Stoma width (μm)	0.31 ± 0.03^c	0.54 ± 0.03^b	0.76 ± 0.05^a
气孔密度 Stoma density (mm^-2)	197.33 ± 18.63^a	43.61 ± 4.51^b	44.21 ± 2.79^b
气孔长/宽 Stoma length-to-width	5.16 ± 0.33^b	6.35 ± 0.41^a	4.55 ± 0.27^b

图3 慈竹不同发育阶段的笋体、竹箨和成熟叶片的叶绿素(平均值±标准差)。I1, 节间伸长期; I2, 节间完全伸长期; I3, 节间成熟期; L, 竹子的成熟叶片; S1, 幼嫩箨鞘; S2, 成熟箨鞘; SL1, 幼嫩箨叶; SL2, 成熟箨叶。不同小写字母表示笋体、竹箨、成熟叶片之间差异显著(p < 0.05)。

Fig. 3 Chlorophyll of shoot body, bamboo sheath and mature branch leaf at different developmental stages of Bambusa emeiensis (mean ± SD). I1, elongating internode; I2, fully elongated internode; I3, mature internode; L, mature branch leaf; S1, young culm sheath; S2, mature culm sheath; SL1, young sheath leaf; SL2, mature sheath leaf. Different lowercase letters indicate significant differences among shoot body, bamboo sheath and mature branch leaf (p < 0.05).

图4 慈竹不同发育阶段的笋体、竹箨和成熟叶片的光合速率、蒸腾速率、气孔导度(平均值±标准差)。I1, 节间伸长期; I2, 节间完全伸长期; I3, 节间成熟期; L, 竹子的成熟叶片; S1, 幼嫩箨鞘; S2, 成熟箨鞘; SL1, 幼嫩箨叶; SL2, 成熟箨叶。不同小写字母表示笋体、竹箨、成熟叶片之间差异显著(p < 0.05)。

Fig. 4 Stomatal conductance (Gs), net photosynthetic rate (Pn), transpiration rate (Tr) of shoot body, bamboo sheath and mature branch leaf at different developmental stages of Bambusa emeiensis (mean ± SD). I1, elongating internode; I2, fully elongated internode; I3, mature internode; L, mature branch leaf; S1, young culm sheath; S2, mature culm sheath; SL1, young sheath leaf; SL2, mature sheath leaf. Different lowercase letters indicate significant differences among shoot body, bamboo sheath and mature branch leaf (p < 0.05).

图5 番红溶液在快速伸长的节间扩散(基部第5节间, Int5)。A, B, 正在伸长节间组织的横切和纵切石蜡切片。A中可见大量的薄壁细胞(PC), B中木质部导管(V)连续。C, H, 0.5%番红溶液处理1 h和4 h时茎秆中番红扩散情况。D-G, 处理1 h时徒手切片中番红扩散情况; I-L, 处理4 h时徒手切片中番红扩散情况. D, E, I, J, 白光下的横切和纵切图; F, G, K, L, 红色滤光片(510-560 nm)下的番红荧光。

Fig. 5 Diffusion of safranine solution in the rapid elongating bamboo shoot (5th internode at base, Int5). A, B, the paraffin section in the elongating internodes. A, a large number of parenchymal cells (PC) are shown, and B shows the continuous xylem vessel (V). C, H, the diffusion of safranine in culm as treated with 0.5% safranine solution for 1 h and 4 h, respectively. D-G, I-L, the diffusion of safranine in the freehand section were showed after 1 h and 4 h treatment, respectively. D, E, I and J, the transverse and longitudinal sections respectively under white light, while F, G, K and L, the corresponding safranine fluorescence under the red filter (510-560 nm) respectively.

图6 竹节中存在“类板状结构” (PLS)。A, B, 茎秆中番红溶液流入幼嫩竹箨。B为A矩形框的纵切徒手切片, 可见番红溶液在竹节处转向竹箨。C, 在竹节部观察到类板状结构(PLS)存在。D, 竹节纵切的连续石蜡切片的组合图, 可见PLS及纵向导管。E, 竹节的横切图, PLS在不同方向存在, 贯穿竹节。F, G, C中①和②处的石蜡切片横切图, 可见PLS中存在大量的导管(V)和韧皮部(P)。图中箭头a和c分别指示箨环(竹箨在竹节的着生处)和秆环(竹节与节间的分隔处), 虚线(不成框)表示竹节位置; 箭头b示节间的纵向导管, d为竹隔。

Fig. 6 A “plate-like structure” (PLS) in the bamboo node. A, B, the safranine solution in the culm flowed into the young bamboo sheath. B was the free-hand longitudinal section of the rectangular frame in A, in which the safranine solution was found turnning into the culm sheath at the node. C, a PLS was showed. D, a serial longitudinal paraffin section of bamboo node, showing the presence of PLS and longitudinal vessel. E, the free-hand transverse section of the node, in which PLS existed in different directions and running through the node. F, G, transverse paraffin sections at ① and ② in C, respectively, there were a large number of vessels (V) and phloem (P) in the PLS. Arrows a and c in A-D pointed the sheath ring (the joint between the culm sheath and the node) and the rod ring (the joint between the bamboo node and the node), respectively. The dashed line (no box) in A-D show the bamboo node; arrow b shows the longitudinal vessels of internodes, and d is the bamboo septum.

图7 离体条件下不同高度竹笋的番红溶液示踪(平均值±标准差)。A, 竹笋各部分的名称。B, 竹笋茎秆的日伸长速度及伸长部分占比。C, D, 竹笋茎秆中番红溶液向上运输的高度及占比。E, F, 将竹笋外层所有竹箨切离后, 茎秆中番红溶液运输高度及占比。伸长部分占比=正在伸长部分/笋体净高; 番红上升比=番红上升高度/笋体净高。

Fig. 7 Tracing of safranine solution of bamboo shoots at different development stages in vitro (mean ± SD). A, Index picture of bamboo shoot. B, Daily elongation rate and elongation ratio of bamboo shoots with different height. C, D, The height and proportion of safranine transported upward respectively in the intact bamboo shoots wrapped by sheaths. E, F, The height and proportion of safranine transported upward respectively in pure shoot body when culm sheaths were separated. Elongation ratio = elongating part/shoot body height; rise ratio of safranine solution = rise height of safranine/shoot body height.

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竹笋期竹箨和笋体的日间蒸腾特性及其对水分运输的影响

Diurnal transpiration of bamboo culm and sheath and their potential effects on water transport during the bamboo shoot stage

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