Anatomical characteristics of resin ducts and its response to extemal stimuli in <i>Pinus yunnanensis</i>

XIONG Hui; LU Xuejia; YANG Wenyun; LI Siguo; GAO Chengjie; LIU Fangyan; LI Kun

doi:10.11833/j.issn.2095-0756.2018.03.012

Volume 35 Issue 3

May 2018

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XIONG Hui, LU Xuejia, YANG Wenyun, LI Siguo, GAO Chengjie, LIU Fangyan, LI Kun. Anatomical characteristics of resin ducts and its response to extemal stimuli in Pinus yunnanensis[J]. Journal of Zhejiang A&F University, 2018, 35(3): 483-489. doi: 10.11833/j.issn.2095-0756.2018.03.012

Citation:

XIONG Hui, LU Xuejia, YANG Wenyun, LI Siguo, GAO Chengjie, LIU Fangyan, LI Kun. Anatomical characteristics of resin ducts and its response to extemal stimuli in Pinus yunnanensis[J]. Journal of Zhejiang A&F University, 2018, 35(3): 483-489. doi: 10.11833/j.issn.2095-0756.2018.03.012

Anatomical characteristics of resin ducts and its response to extemal stimuli in Pinus yunnanensis

doi: 10.11833/j.issn.2095-0756.2018.03.012

1.
Research Institute of Resource Insects, Chinese Academy of Forestry, Kunming 650224, Yunnan, China
2.
Meiziqing Forest Farm of Shuangbai County, Shuangbai 675106, Yunnan, China

Received Date: 2017-05-27
Rev Recd Date: 2017-08-05
Publish Date: 2018-06-20

Abstract

Oleoresin is a secondary metabolite with a high economic and ecological value, and it is mainly stored in the resin channel system of needles and trunk xylem and phloem of the genus Pinus. Study on the relationship between oleoresin yield and their anatomical characteristics is conducive to selective breeding of superior tree with high-yielding oleoresin of Pinus yunnanensis and establishment of directional cultivation technology system of high-yielding forest. The relationship between oleoresin and the anatomical characteristics of needles and trunks was studied, and the differences in the anatomical characteristics of the needles and trunks of high-and low-yielding trees were compared and analyzed. The results showed that, the number of resin canals and secrete cells in the needles and trunks of high-yielding oleoresin pines were significantly higher than those of low-yielding oleoresin pines. With the increase of the number of resin canals in the needles and trunks, the oleoresin yield was also increased(P < 0.01). In addition, the loading of the number of resin canal in xylem was the largest in the principal component analysis. The result implied that the number of resin canal in xylem could be proposed as an important indicator for breeding high-yielding oleoresin P. yunnanensis.
- forest tree breeding,
- Pinus yunnanensis,
- oleoresin yield,
- anatomical characteristics

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沈阳化工大学材料科学与工程学院沈阳 110142

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松脂是由松属Pinus树种中分泌细胞分泌而成的一种主要由萜烯类化合物构成的无色透明状次生代谢产物, 储藏在松树针叶以及枝和茎的木质部与韧皮部的树脂道系统内^[1-6]。松脂可分离出松节油和松香, 具有很高的经济价值和生态价值。松节油具有很强的同分异构能力, 是一种良好的溶剂, 广泛应用于油漆、冶金、纺织等工业领域, 此外可用于生产一些药物, 如驱蛔脑、冰片、樟脑等, 以及聚萜烯树脂等多种树脂。松香广泛用于印刷油墨、绝缘材料、造纸施胶、涂料、油漆、合成橡胶、肥皂、人造甘性油、软化剂和干燥剂等领域。如今, 人们还对松香进行了改性, 生产出聚合松香、歧化松香和氢化松香等一系列产品, 使松香的价值得到了进一步提高^[7-8]。云南松Pinus yunnanensis是松属的一种常绿乔木, 以滇中高原为中心, 向四周分布, 整个分布区呈现不规则的多角形状, 其分布区面积占云南省森林面积的70%。成林主要分布在海拔1 000~3 200 m的地区, 是西南地区主要的采脂树种^[9]。云南松单株之间泌脂量差异明显, 单株年产脂量最高可达11.00 kg, 而最低的甚至小于1.00 kg, 直接影响了人们对云南松采脂利用的经济效益。研究表明:高产脂马尾松Pinus massoniana针叶树脂道个数明显高于普通产脂量类型, 针叶树脂道数目与矫正产脂量、日产脂量和10 cm侧沟产脂量均存在显著相关。马尾松针叶树脂道个数越多, 其产脂力越高^[10]。此外, 分泌细胞个数与思茅松Pinus kesiya var.langbianensis的产脂力有密切关系, 分泌细胞个数越多, 其产脂力越高^[11]。关于云南松产脂量与针叶及树干解剖学之间关系的研究较少。鉴于此, 本研究以不同产脂量的云南松为对象, 研究云南松的解剖学特征及外界刺激对它产生的影响, 旨在为选育云南松高产脂优树和建立高产脂林定向培育技术体系提供依据。

1. 研究区概况

研究区位于滇中高原楚雄州双柏县(24°19′~24°22′N, 101°34′~101°37′E), 该区海拔为1 600~1 800 m, 气候垂直变化比较明显, 属于北亚热带季风气候。年平均气温为15.5 ℃, 极端最高气温为42.0 ℃, 极端最低气温为-8.4 ℃。雨热同季, 干湿分明, 年平均降水量约为900 mm, 日照充足, 年平均日照时间为2 355 h。土壤为偏酸性, 属于含钾、磷元素比较丰富的山地红壤。研究区以云南松纯林为主, 其中伴生了少量的麻栎Quercus acutissima, 木荷Schima superba, 滇青冈Cyclobalanopsis glaucoides, 锐齿槲栎Q.aliena var.acuteserrata, 高山栲Castanopsis delavayi和旱冬瓜Alnus nepalensis等。

2. 材料与方法

2.1. 采脂

在双柏县太和江林场, 根据当地脂农提供的该区域云南松多年的产脂情况以及现场实地勘察结果, 选择了松脂产量相对稳定且较高的云南松林地, 运用“S”型布点, 在同一等高线附近, 选择胸径大于20.0 cm并且已采2 a以上的生长状况基本一致的云南松60株, 用铁皮标牌依次给60株云南松编号。同时, 从2015年3月初至10月底对所选60株云南松进行采脂。采脂标准为:采脂面选择云南松树干无结疤且向阳面, 采割位置距离地面2.2 m, 先割中沟且垂直于地面, 再割侧沟且夹角为70°左右, 采割深度到木质部, 但不超过0.5 cm, 采割面的负荷率不能超过50%并且长度为20.0 cm^[12-14], 隔2 d采割1次, 每个月统计60株云南松的产脂量, 共统计8次, 以每株云南松8个月的总产脂量作为单株年产脂量。

2.2. 试验样品采集

2.2.1. 解剖样品取样

根据60株云南松产脂分组情况(组数为12组, 组距为1.00 kg), 各组取1/2该组的株数, 若所得是小数, 则去除小数点后面取整数株, 若该组为1, 则取1株, 若为0, 则不取。共取30株, 用人工攀树和高枝剪相结合的方法在云南松的树冠中部向阳方向采取1年生针叶, 共采取针叶10束·株^-1, 洗净, 并用吸水纸吸干, 用剪刀取5.0 cm针叶中部放入固定液中(20.0 g·kg^-1多聚甲醛和12.5 g·kg^-1戊二醛, 50 mmol·L^-11, 4哌嗪二乙烷磺酸, pH 7.2)并编号, 共30个样, 带回实验室制作切片(试验中所用固定液均与此相同)。在每株树距离地面3.5 m高的位置, 去掉树皮(软木层), 记号笔画出2.0 cm × 2.0 cm的矩形, 用1.0 cm的凿子, 凿取2.0 cm × 2.0 cm的样品(包括韧皮部、形成层和木质部), 分别放入固定液中, 共30个样, 带回实验室制作切片, 用硅胶塞封住取样所造成的伤口。

2.2.2. 茉莉酮酸甲酯实验取样

根据年产量大于8.41 kg为高产, 小于3.35 kg为低产的标准, 在30株云南松中选出高产和低产各5株, 根据胸径位置的周长, 将各株云南松树干均分成3个区域, 依次编号A, B, C。第1次取样:A区域为第1次取样区域, 在A区域距离地面3.5 m高的位置, 去掉树皮, 凿取2.0 cm × 2.0 cm的样品, 1块·株^-1, 共10块, 分别放入固定液中(2.2.1中已经取过样品)。在与A区域取样同一高度, 去掉树皮(软木层), 将配好的100 mmol·L^-1茉莉酮酸甲酯溶液(茉莉酮酸甲酯+1.0 g·kg^-1吐温20)喷在B区域, 用棉签涂抹该区域, 防止液体流出, 重复上述操作, 使该区域保湿5 min。在与A区域和B区域同样的处理高度, 用1.0 g·kg^-1吐温20处理C区域^[15-16]。第2次取样:处理后8 d, 在第1次取样及试剂处理的区域上方5.0~20.0 cm处分别取样, 取样方法及样品处理同第1次取样。第3次取样:处理后35 d, 在第1次取样及试剂处理的区域下方5.0~20.0 cm处分别取样, 取样方法及样品处理同第1次取样。

2.3. 试验样品处理

将带回实验室的样品进行修整, 针叶切成2.0 cm, 木质部和韧皮部切成1.0 cm × 1.0 cm, 用蒸馏水清洗修正后的样品, 去除样品上多余的固定液, 并用石蜡切片法将样品制成12.0 μm的切片, 经番红固绿染色后, 中性树胶封片, 然后用显微镜观察样品树脂道大小及分泌细胞个数

2.4. 数据处理与分析方法

用SPSS 17.0统计软件, 运用Pearson相关系数评价30株云南松产脂量与其针叶和树干解剖特征之间的相关关系, 同时, 对所选高、低产脂各5株云南松针叶和树干解剖特征运用单因素方差分析(one-way ANOVA)比较差异显著性, 并用Excel 2007制图。

4. 结论与讨论

树脂道是广泛存在于松树针叶和树干中的一种解剖器官, 主要包括本构性树脂道和诱导性树脂道。本构性树脂道是松树固有的一种结构, 而诱导性树脂道则是植物受到外界昆虫入侵或机械损伤时所形成的一种防御结构。本研究表明:高、低产脂量云南松针叶树脂道个数、韧皮部和木质部树脂道个数均表现出差异极显著(P＜0.01), 而且, 随着这些树脂道的增加, 云南松产脂量也不断增加(P＜0.01), 这与朱秋云等^[10]、王海林等^[17]的研究结果相似。可能是因为松树自身的防御机制与树脂道有关, 而松脂是松树分泌的一种次生代谢产物, 存储在树脂道中, 当松树受到机械损伤或者昆虫入侵, 存储在本构性树脂道中本构性树脂迅速溢出, 杀死昆虫及病菌并封闭伤口, 同时诱导形成大量诱导性树脂道及诱导性松脂^[18-19]。

分泌细胞也是植物解剖结构的一个关键, 它参与了植物体内多种生理生化反应。本研究表明:高产脂云南松针叶及树干分泌细胞极显著高于低产脂云南松(P＜0.01), 而且, 随着这些分泌细胞个数的增加, 云南松产脂量也不断增加(P＜0.01)。这与穆茹等^[11]所得结论相同。松脂是由分泌细胞分泌的^[18], 松树在受到外界刺激时, 存储在本构性树脂道中的松脂就会大量流出, 同时形成大量新的分泌细胞, 分泌更多的松脂^[19-20]。

主成分分析发现:木质部树脂道个数可作为选取高产脂量云南松的指标。本研究还发现:机械损伤和茉莉酮酸甲酯处理能增加云南松树脂道个数和分泌细胞个数, 并且, 低产脂云南松韧皮部树脂道个数、木质部树脂道个数和分泌细胞个数增加的量比高产脂云南松多, 但是否能提高云南松产脂量有待进一步研究。

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产脂类型	树脂道数/个	树脂道平均面积/μm²	分泌细胞数/个
高产月旨	9.75 ± 0.50 aA	83.54 ± 28.74 aA	78.75 ± 1.26 aA
低产脂	2.50 ± 0.58 bB	65.51 ± 18.29 aA	14.50 ± 3.51 bB
说明:数值为平均值±标准差，同一指标不同小写字母表示差异显著(P < 0.05)，不同大写字母表示差异极显著(P < 0.01)

产脂类型	韧皮部树脂道数/(个·cm^-2)	韧皮部树脂道直径/mm	木质部树脂道数/(个·cm^-2)	木质部树脂道面积/μm²/	分泌细胞数/个
高产脂	100.63 ± 3.75 aA	1.91 ± 0.56 aA	88.13 ± 10.28 aA	140.66 ± 3.97 aA	317.25 ± 7.02 aA
低产脂	56.25 ± 6.29 bA	1.87 ± 0.47 aA	13.13 ± 2.39 bA	148.00 ± 8.30 aA	31.50 ± 5.75 bA
说明:数值为平均值±标准差，同一指标不同小写字母表示差异显著(P < 0.05)，不同大写字母表示差异极显著(P < 0.01)

取样次序	韧皮部树脂道数/(个·cm^-2)		韧皮部树脂道直径/mm		木质部树脂道数/(个·cm^-2)		木质部树脂道面积/μm²		分泌细胞数/个
取样次序	高产脂	低产脂	高产脂	低产脂	高产脂	低产脂	高产脂	低产脂	高产脂	低产脂
1	P 100.63±3.75 A 110.88±1.88 a	56.25±6.29 66.63±1.63 a	1.91±0.56 2.15±0.18 a	1.87±0.46 2.19±0.34 a	88.13±10.28 92.98±5.90 a	13.13±2.39 18.15±2.53 a	140.66±3.97 l45.68±3.64 a	148.00±8.30 l53.05±8.56 a	317.25±7.02 320.26±2.65 a	31.50±5.75 34.95±1.05 a
2	B 109.05±1.08 a C 100.65±2.88b A 117.13±1.20 a	68.35±2.08 a 56.20±4.95b 75.90±3.38 a	2.09±0.21 a 1.92±0.52 a 2.26±0.24 a	2.10±0.25 a 1.86±0.18b 2.32±0.23 a	91.70±3.95 a 88.13±2.65b 95.35±5.30 a	17.58±1.98 a 13.13±2.13b 21.73±1.00 a	140.59±4.12 a 140.66±3.59 a 146.67±3.12 a	148.04±7.98 a 148.00±9.12 a 146.67±3.12 a	319.65±2.84 a 317.24±3.56b 322.29±4.56 a	33.89±0.27 a 31.49±0.48b 37.01±1.25 a
3	B 114.88±0.65 a C 100.60±3.13 b	72.83±2.45 a 56.23±5.33 b	2.19±0.14 a 1..91±0.63 a	2.25±0.19 a 1.87±0.42 a	94.23±4.95 a 88.10±2.23 b	20.15±0.63 a 13.13±4.13 b	140.66±4.15 a 140.65±3.76 a	148.02±7.69 a 148.03±8.17 a	320.05±3.15 a 317.25±4.12 b	34.59±0.95 a 31.50±2.01 b
说明：数值为平均值±标准差，P表示未做任何处理，A，B和C分别表示机械损伤、茉莉酮酸甲酯处理和l.0g·kg^-1吐温20处理。每次取样不同处理间字母不同表示差异显著(P < 0.05)

成分	提取量	特征根	贡献率/G	累积贡献率/G	载荷
针叶树脂道个数	0.957	4.832	96.641	96.641	0.978
针叶分泌细胞个数	0.987	0.125	2.498	99.139	0.994
韧皮部树脂道个数	0.972	0.030	0.598	99.737	0.986
木质部树脂道个数	0.990	0.008	0.166	99.903	0.995
树干分泌细胞个数	0.926	0.097	0.097	100.000	0.962

Anatomical characteristics of resin ducts and its response to extemal stimuli in Pinus yunnanensis

doi: 10.11833/j.issn.2095-0756.2018.03.012