Anatomical Observation on the Tortuousness of Ziziphus jujuba var. tortuosa Branches

doi:10.16420/j.issn.0513-353x.2020-0561

Acta Horticulturae Sinica ›› 2021, Vol. 48 ›› Issue (9): 1653-1664.doi: 10.16420/j.issn.0513-353x.2020-0561

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Anatomical Observation on the Tortuousness of Ziziphus jujuba var. tortuosa Branches

GUO Xuemin^*(), WANG Xinrui, WANG Yingying, LI Zheng, MIAO Ningning, WANG Zhaojun

College of Oceanic Resources and Environment,Hebei Normal University of Science & Technology,Changli,Hebei 066600,China

Received:2021-06-18 Revised:2021-08-20 Online:2021-09-25 Published:2021-09-30
Contact: GUO Xuemin E-mail:xueminguo@126.com

Abstract

Abstract:

Taking the twisted tortuous branch segment and twisted straight branch segment of annual branches in Ziziphus jujuba Mill. var. tortuosa as experimental materials,the morphological,anatomical and vessel characteristics of its branches were observed and compared by xylotomy and wood segregation techniques. The differences of anatomical structures between twisted tortuous branch segment and twisted straight branch segment were mainly manifested in the number,shape and distribution of abnormal secondary structures. Compared with the twisted straight branch segment,the number of auxillary bundles in twisted tortuous branch segments was more,and there were three auxillary bundles adjacent to each other,which were relatively concentrated in the inner bend of the segments;The number of interxylary phloem was about the same as that of the twisted straight branch segment,and its end expanded,which was different from the slightly curved elliptic columnar shape of the twisted straight branch segment,and relatively concentrated in the outer bend of the segments;The number of medullary vascular bundles was slightly reduced. At the cellular level,the vessel elements at the inner and outer bends of the twisted tortuous branch segments of the variety were all reticulate vessels with single perforation and various shapes. Among them,the number of vessel elements with tails at both ends and sloped at both ends at the inner bend were 28.0% and 20.2% less than those at the outer bend,respectively. The number of vessels without tails at either ends,sloped at one end and horizontal at both ends were 17.9%,375.0% and 200.0% more,respectively. The number of vessels with one tail at one end was the same. The end wall slope of end wall was 10.1% larger than that at the outer bend,and the length of vessel element was 16.4% shorter than that at the outer bend,while the diameter was 18.6% smaller than that at the outer bend. The hypothesis of “abnormal structure-spatial difference-coordination”was put forward. The tortuous branch traits of Z. jujuba var. tortuosa may be the result of the unbalanced growth rates and forces at different locations of the branches caused by the spatial difference and coordination of the number,the mechanical tension,the transport and utilization of the assimilated products of the three abnormal secondary structures,namely the auxillary bundle,interxylary phloem and medullary bundle.

Key words: Ziziphus jujuba, tortuous branch, abnormal secondary structure, vessel element

CLC Number:

S665.1

GUO Xuemin, WANG Xinrui, WANG Yingying, LI Zheng, MIAO Ningning, WANG Zhaojun. Anatomical Observation on the Tortuousness of Ziziphus jujuba var. tortuosa Branches[J]. Acta Horticulturae Sinica, 2021, 48(9): 1653-1664.

Figures/Tables 6

Fig. 1 The branch shape of Ziziphus jujuba var. tortuosa TB：Twisted tortuous branch segment;TS：Twisted straight branch segment;IS;The inner bend of twisted tortuous branch segment;OS：The outer bend of twisted tortuous branch segment.

Fig. 2 Comparison of anatomical structures of bark between twisted straight and twisted tortuous branch segments of Ziziphus jujuba var. tortuosa branches 1. Twisted straight branch segment;2. Twisted tortuous branch segment. a1. Complete branch transection,showing distribution of five scattered auxillary bundles（AB）,two interxylary phloems（IP）and four medullary bundles（MB）;a2. Transection of the branch,showing the incision（arrow）marking the inner bend of twisted tortuous branch segment,and that nine of the ten auxillary bundles which consisted of two adjacent auxillary bundles and four separate auxillary bundles,in which one adjacent auxillary bundle contained three auxillary bundles,and the other contained two auxillary bundles,were relatively concentrated on the inner bend of the branch,while four of the five interxylary phloems were relatively concentrated on the outer bend of the branch;b. The bark,showing epidermis（Ep）,periderm（Pe）,cortex（Co）,secondary phloem fiber band（SB）and secondary phloem（SP）;c. Longitudinal section of the bark,showing the structure of the closed lenticel （Le）,two parallel secondary phloem fiber bands（SB）and two secretory cavities（SC）;d1. Detailed view of one off-center auxillary bundle transection formed by the fission of secondary xylem（SX）,showing the asymmetric structure of its xylem and lateral secondary phloem fiber band;d2. Transection of an in-center auxillary bundle originated from peripheral parenchyma cells;e1. Two in-center auxillarybundles（AB）originated from peripheral parenchyma cells and one developing and in-center auxillary bundle（circle）originated from peripheral parenchyma cells,showing the secondary phloem fiber band on the inside of auxillary bundles;e2. Transection of a off-center auxillary bundle formed by the fission of secondary xylem,showing the pattern of its connection with secondary xylem（arrow）and its outer arc-shaped secondary phloem fiber band;f1. Longitudinal section of one auxillary bundle（AB）originated from peripheral parenchyma cells,the arrow showing some secondary xylem cells arranged perpendicular to the long axis of the branch. h1. Longitudinal section through the end of one auxillary bundle,showing a spherical end of an auxillary bundle consisting of irregularly arranged secondary xylem cells;i1. Transection of two adjacent auxillary bundles;j1. Detailed view of transection of one off-center auxillary bundle formed by the fission of secondary xylem,showing its connection with secondary xylem;f2. Transection of three adjacent auxillary bundles connected together,showing one auxillary bundle（Left）originated from peripheralparenchyma cells and two auxillary bundles formed by the fission of secondary xylem,the pattern of their interconnection and the distribution of secondary phloem fiber band;g2. Longitudinal section of one auxillary bundle originated from peripheral parenchyma cells,showing its fusiform shape and secondary phloem band located in the inner side of the auxillary bundle;h2. Longitudinal section of one auxillary bundle fiberformed by the fission of secondary xylem,showing its oval shape,secondary phloem fiber band located outside the auxillary bundle and two secretory cavities;k1. Tangential section of one branch, showing two auxillary bundles（AB）originated from peripheral parenchyma cells（Independent of the secondary xylem）,in which the secondary phloem fiber band（SB）was on its inner side,and one auxillary bundle（AB）（Left）formed by the fission of secondary xylem（Associated with the secondary xylem）,in which the secondary phloem fiber band（SB）was outside it（double arrows）;i2,j2. Section of one auxillary bundle connecting with secondary xylem,Longitudinal showing two secondary phloem fiber bands located outside the auxillary bundle. PX. Primary xylem;Sc. Sclereid;Pi. Pith.

Fig. 3 Comparison of anatomical structures of secondary xylem and pith between twisted straight and twisted tortuous branch segments of Z. jujuba var. tortuosa branches 1. Twisted straight branch segment;2. Twisted tortuous branch segment. a1. Transection of secondary xylem,showing vessels（V）and uniserate rays （UR）;b1. Tangential section of secondary xylem,showing reticulate vessel and the height of uniserate ray;c1. Transection of the branch, showing the distribution of interxylary phloems（IP）and secondary phloem fiber band outside secondary xylem;d1. Detailed view of transaction of the outer part of an interxylary phloem,showing composition of the phloem;e1. Detailed view of pith end of interxylary phloem,showing unidirectional cambium;f1. Tangential section of secondary xylem,showing the oval shape of the interxylary phloem transection;g1. Transection of the pith;h1. Longitudinal section of pith;i2. Detailed view of figure a2 in fig.2,showing the distributions of four interxylary phloems and their shapes of enlarged ends,and the shape of one medullary bundle（MB）;j1. Detailed view of one medullary bundle transection,showing its nearly symmetrical shape;k1. Detailed view of one elliptical medullary bundle transection,showing its asymmetric shape;l1. Detailed view of longitudinal section through one medullary bundle,showing central phloem tissue and peripheral secondary xylem. AB：Auxillary bundles;SB：Secondary phloem fiber band;PX：Primary xylem;Pi：Pith.

Fig. 4 Vessel element shapes in twisted tortuous branch segment of Ziziphus jujuba var. tortuosa branches A. Both end walls are horizontal,and tails in both end walls;B. Both end walls are slope,and tail in one end wall;C. Both end walls are slope,and no tail in either end wall;D. One end wall is horizontal,and tail in one end wall;E. Both end walls are near horizontal,and no tail in either end walls;F. Both end walls are slope,and tails in both end walls.

Table 1 Size and frequency of tail types of vessel elements in twisted tortuous branch segment of Z. jujuba var. tortuosa

部位 Position	大小/µm Size		尾类型 Tail type
部位 Position	长度 Length	直径 Diameter	两端有尾 Both tail	一端有尾 One tail	两端无尾 No tail
内弯侧 Inner bend	259.3 ± 79.1	39.8 ± 11.5	18	36	46
外弯侧 Outer bend	310.3 ± 31.2^**	48.9 ± 11.4^*	25	36	39

Table 2 Frequency of types of end wall slope of vessel elements in twisted tortuous branch segment of Z. jujuba var. tortuosa

材料 Material	端壁类型比例/% End wall type			端壁倾角/° End wall slope
材料 Material	两端倾斜 Both end walls are slope	一端倾斜 One end wall is slope	两端水平 Both end walls are horizontal	端壁倾角/° End wall slope
内弯处The inner bend	75	19	6	53.2 ± 13.9^**
外弯处The outer bend	94	4	2	48.3 ± 11.9

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Anatomical Observation on the Tortuousness of Ziziphus jujuba var. tortuosa Branches

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