林业科学 ›› 2023, Vol. 59 ›› Issue (8): 40-51.doi: 10.11707/j.1001-7488.LYKX20220802
徐薪璐1,2,3,蔡鸥1,2,3,赵婉琪1,2,3,吕卓1,2,3,姚文静1,2,李龙1,2,林树燕1,2,*()
收稿日期:
2022-11-18
出版日期:
2023-08-25
发布日期:
2023-10-16
通讯作者:
林树燕
E-mail:lrx@njfu.com.cn
基金资助:
Xinlu Xu1,2,3,Ou Cai1,2,3,Wanqi Zhao1,2,3,Zhuo Lü1,2,3,Wenjing Yao1,2,Long Li1,2,Shuyan Lin1,2,*()
Received:
2022-11-18
Online:
2023-08-25
Published:
2023-10-16
Contact:
Shuyan Lin
E-mail:lrx@njfu.com.cn
摘要:
目的: 探究靓竹彩叶形成过程中细胞结构和光合色素的变化,以及不同分区的叶色与光合色素、细胞结构的关系,为明确靓竹叶片发育过程中条纹叶色的形成机制提供参考。方法: 以靓竹不同发育时期的叶片为材料,通过扫描电镜观察不同发育时期叶片内部显微结构的变化,并将叶片按照叶色分为黄区和绿区,制作徒手切片,置于光学显微镜下观察以比较叶片黄区和绿区的内部结构。同时测定每个时期的光合色素和叶绿素合成前体物质含量。结果: 不同发育过程中靓竹各叶片的显微结构、超微结构、光合色素含量、叶绿素合成前体物质含量差异显著,成熟功能叶不同分区的叶片结构和光合色素含量之间也存在显著差异。1)叶片发育过程中表皮细胞、叶肉细胞逐渐增大,叶脉面积增大。2)发育过程中绿区叶肉细胞内叶绿体发育正常,内部结构完整。横切面细胞中的叶绿体数量增加,基粒片层数逐渐增加,内部排列趋于紧密;黄区叶肉细胞分化程度低,指状臂细胞层数少,细胞内空腔大,叶绿体发育异常。3)叶片黄区和绿区之间叶片表皮结构差异不显著,但黄区和绿区叶肉细胞叶绿体分布位置差异明显,绿区叶肉细胞中均有叶绿体分布,而黄区叶绿体仅分布于靠近叶片下表皮的位置。4)光合色素含量随叶片发育过程逐渐累积,绿区的各色素含量显著高于黄区。5)尿卟啉原III和粪卟啉原III是靓竹叶片发育过程中叶绿素合成途径中的关键物质,伴随叶片条纹的出现,尿卟啉III向粪卟啉原III的转化增加,但粪卟啉原III向原卟啉IX的转化减少。结论: 靓竹叶片发育过程中,叶片厚度逐渐增大,叶片表皮细胞、叶肉细胞、叶脉均不断增大,光合色素含量逐渐积累。绿区叶片随发育程度叶绿体数量逐渐增加,叶绿素含量积累。但黄区叶片叶肉细胞空腔大且叶绿体发育异常,指状臂细胞层数少。靓竹条纹叶色的出现与指状臂细胞分化程度低、叶绿体发育异常以及叶绿素合成受阻有关。
中图分类号:
徐薪璐,蔡鸥,赵婉琪,吕卓,姚文静,李龙,林树燕. 靓竹彩叶形成的细胞学以及光合色素[J]. 林业科学, 2023, 59(8): 40-51.
Xinlu Xu,Ou Cai,Wanqi Zhao,Zhuo Lü,Wenjing Yao,Long Li,Shuyan Lin. Changes in Cytology and Photosynthetic Pigments during the Formation of Colored Leaves of Sasaella glabra ‘Albostriata’[J]. Scientia Silvae Sinicae, 2023, 59(8): 40-51.
表1
靓竹叶片不同发育时期特征描述"
时期Stage | 特征描述Description of leaf characteristics |
S1 | 位于靓竹枝条顶部且被叶鞘包裹,未露出叶鞘, 卷曲,嫩白色 Located at the top of branch and wrapped in a leaf sheath, without exposing the leaf sheath, curled, tender white |
S2 | 位于枝条顶部且刚伸出叶鞘0.5 cm以内,卷曲,嫩黄色叶片出现浅绿色条纹 Located at the top of the branch and just protruding within 0.5 cm of the leaf sheath, curled, with light green stripes on tender yellow leaves |
S3 | 位于靓竹枝条顶部,伸出叶鞘,舒展中,具黄绿相间条纹 Located at the top of branch, protruding from the leaf sheath, spreading out with alternating yellow and green stripes |
S4 | 功能叶,叶片完全展开,黄绿相间条纹明显 Functional leaves, fully unfolded, with distinct yellow green stripes |
表2
靓竹叶片不同生长阶段显微结构指标(一)①"
时期 Stage | 叶片厚度 Leaf thickness/μm | 上表皮厚度 Upper epidermis thickness/μm | 下表皮厚度 Lower epidermis thickness/μm | 泡状细胞 Bulliform cell length/μm | 梅花状细胞 Rosette cell thickness/μm | 指状臂细胞 Arm cell thickness/μm | 梭形细胞 空隙长度 Fusoid cell thickness/μm | |
S1 | 88.40±6.96c | 8.63±1.67c | 8.89±1.51c | 29.55±10.39e | 未分化Undifferentiation | 未分化Undifferentiation | 未分化Undifferentiation | |
S2 | 97.78±5.07b | 9.75±1.66b | 9.47±1.15c | 35.94±11.40d | 未分化Undifferentiation | 未分化Undifferentiation | 未分化undifferentiation | |
S3 | 100.69±6.92b | 9.97±1.63b | 10.41±1.75bc | 55.09±14.29c | 12.50±2.41b | 15.40±2.91a | 58.14±10.86a | |
S4 | 黄区Yellow stripe | 103.44±17.07b | 11.24±1.321a | 13.82±2.53a | 64.03±11.69b | 13.44±2.50a | 16.39±3.64a | 63.43±9.83a |
绿区 Green stripe | 108.25±8.46a | 11.80±1.51a | 14.02±1.83a | 85.05±12.48a | 13.65±2.62a | 16.36±3.10a | 60.74±9.57a |
表3
靓竹叶片不同生长阶段显微结构指标(二)①"
时期 Stage | 主脉面积 Area of main vascular/μm2 | 1级脉面积 Area of first vascular/μm2 | 2级脉面积 Area of second vascular/μm2 | |
S1 | 3 000.83±484.59d | 1 639.58±435.65e | 453.12±109.22c | |
S2 | 3 776.53±506.92c | 2 711.88±519.29d | 555.10±132.80b | |
S3 | 5 152.21±452.31b | 3 877.31±940.31c | 617.41±133.23b | |
S4 | 黄区 Yellow stripe | 9 076.42±1425.09a | 4 776.94±493.83b | 775.68±279.24a |
绿区 Gre en stripe | 9 155.99±1260.08a | 5 298.49±661.75a | 806.14±198.89a |
图3
靓竹不同发育时期叶片超微结构观察 a1–4代表不同放大倍数下花叶S1时期叶片细胞结构;b1–2、c1–2、d1–2分别为不同放大倍数下花叶S2–4时期绿区叶片细胞结构;b3–4、c3–4、d3–4分别为不同放大倍数下花叶S2–4时期黄区叶片细胞结构; CN:细胞核; Ch: 叶绿体; G: 基粒; S: 淀粉粒; OG: 嗜饿颗粒; CW:细胞壁;P:质体;MC:线粒体;GA:高尔基体。a1–4 represents the leaf cell structure at S1 stage under different magnification; b1–2, c1–2 and d1–2 were the cell structures of green district leaves at S2–4 stage under different magnification; b3–4, c3–4 and d3–4 are the cell structures of yellow area leaves at different magnification at S2–4 stage of mosaic respectively.CN: Cell nucleus; Ch: Chloroplast; G: Granum; S: Starch grain; OG: Osmiophilic granule; CW: Cell wall; P: Plastid; MC: mitochondrion; GA: Golgi apparatus."
表4
靓竹不同时期叶片光合色素含量变化①"
时期Stage | 叶绿素a Chlorophyll a/( mg·g?1) | 叶绿素b Chlorophyll b/( mg·g?1) | 叶绿素 Chlorophyll/( mg·g?1) | 类胡萝卜素 Carotenoids/( mg·g?1) |
S1 | 0.297±0.037c | 0.074±0.032b | 0.370±0.037c | 0.236±0.016c |
S2 | 0.364±0.213c | 0.198±0.090b | 0.562±0.228c | 0.327±0.042c |
S3 | 9.441±0.767b | 2.169±0.496a | 11.610±1.23b | 3.206±0.188b |
S4 | 11.941±2.497a | 2.202±0.471a | 14.143±2.961a | 4.282±0.825a |
表5
靓竹S4时期叶片光合色素含量分区对比①"
表型Phenotype | 叶绿素a Chlorophyll a/( mg·g?1) | 叶绿素b Chlorophyll b/( mg·g?1) | 叶绿素 Chlorophyll/( mg·g?1) | 类胡萝卜素 Carotenoids/( mg·g?1) | 叶绿素/类胡萝卜素 Chlorophyll/Carotenoids |
S4 | 14.186±2.608c | 3.623±1.202b | 17.809±3.806c | 6.247±0.731b | 3.098±0.141a |
YS | 2.169±0.431d | 1.250±0.617c | 3.419±1.045d | 1.158±0.174c | 2.952±0.399c |
GS | 24.977±2.681b | 5.407±0.971b | 30.384±3.648b | 8.943±0.664a | 3.542±0.153b |
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