Effects of storage substances and endogenous hormones on seed germination and seedling formation of Quercus variabilis
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摘要:目的
分析栓皮栎大、中、小粒种子的萌发出苗特性，解析种子贮藏物质和内源激素影响其萌发出苗特性的生理机制，从种子规格角度阐明栓皮栎的更新策略。
方法称量采集的5 000粒种子的单粒质量，聚类分成大粒（Ⅰ）、中粒（Ⅱ）和小粒（Ⅲ）共3个规格等级，后于人工气候培养箱育种并观测萌发出苗动态。采用酶解法测定种子可溶性糖与淀粉含量，考马斯亮蓝G-250染色法测定可溶性蛋白含量，高效液相色谱法测定赤霉素（GA3）、脱落酸（ABA）、吲哚乙酸（IAA）含量，并利用主成分分析法探究影响种子萌发与出苗的主要因素。
结果（1）栓皮栎Ⅰ、Ⅱ、Ⅲ级种子数量所占比例分别为23.82%、42.78%、33.40%。不同规格种子的淀粉、可溶性蛋白、ABA含量均值存在显著差异，Ⅰ级种子的淀粉含量均值分别是Ⅱ、Ⅲ级种子的2.02、2.37倍；Ⅰ 级种子的可溶性蛋白含量均值分别是Ⅱ、Ⅲ 级种子的1.05、1.87倍，Ⅰ 级种子ABA的含量分别是Ⅱ、Ⅲ 级种子的0.86、0.60倍。（2）种子萌发出苗试验结果显示：Ⅰ、Ⅱ、Ⅲ 级种子累计萌发率分别为91%、84%、77%，累计出苗率分别为90%、81%、70%，萌发未出苗的种子比例分别占1%、3%、9%，萌发至出苗结束所需天数分别为17、18、23 d。（3）主成分分析结果显示：贮藏物质与植物激素对种子萌发出苗均具有一定影响，以ABA与可溶性蛋白含量影响最为显著。
结论栓皮栎规格大的种子所占比例小，萌发和出苗速度快，萌发和出苗率高，规格小的种子则相反，资源分配与萌发特性存在权衡；不同规格大小的种子萌发出苗之间的差异受到贮藏物质以及植物激素的影响。因此，人工培育苗木时，先将种子分级后再播种可提高苗木整齐性，降低挑苗、分级育苗的人工成本。
Abstract:ObjectiveThis paper aims to analyze the germination and emergence characteristics of large, medium and small seeds of Quercus variabilis, and elucidates the physiological mechanisms, by which seed storage substances and endogenous hormones affect its germination and emergence characteristics. From the perspective of seed specifications, the regeneration strategy of Q. variabilis was clarified.
MethodThe single-seed mass of 5 000 collected seeds was measured, and classified into three size grades, i.e. large (Ⅰ), medium (Ⅱ), and small (Ⅲ). Then they were subjected to artificial climate cultivation in a growth chamber to observe the germination and seedling dynamics. The soluble sugar and starch content, soluble protein content, content of gibberellin (GA3), abscisic acid (ABA), and indoleacetic acid (IAA) were measured using enzymatic, coomassie brilliant blue G-250 staining, high-performance liquid chromatography (HPLC) methods, respectively. Principal component analysis was used to explore the main factors influencing seed germination and seedling emergence.
Result(1) The number of seeds of Q. variabilis Ⅰ, Ⅱ and Ⅲ accounted for 23.82%, 42.78% and 33.40%, respectively. There were significant differences in the average content of starch, soluble protein, and ABA among three size grades. The average starch content of Ⅰ seeds was 2.02 and 2.37 times of Ⅱ and Ⅲ seeds, the average soluble protein content of Ⅰ seeds was 1.05 and 1.87 times of Ⅱ and Ⅲ seeds, and the ABA content in Ⅰ seeds was 0.86 and 0.60 times of Ⅱ and Ⅲ seeds, respectively. (2) Seed germination and emergence experiment showed that the cumulative germination percents of Ⅰ, Ⅱ and Ⅲ seeds were 91%, 84% and 77%, and the cumulative emergence percents were 90%, 81% and 70%, respectively. The proportion of germination but not emergence accounted for 1%, 3% and 9%, respectively. The days from germination to the end of emergence for Ⅰ, Ⅱ and Ⅲ were 17, 18 and 23 d, respectively. (3) Principal component analysis showed that seed germination and seedling emergence were affected not only by seed reserves, but also by plant hormones, especially the content of ABA and soluble protein.
ConclusionThe proportion of large-sized seeds in Q. variabilis is small, with fast germination and emergence rates, while small-sized seeds are the opposite. There is a trade-off between resource allocation and germination characteristics. The differences in germination and emergence of seeds of different sizes are influenced by storage substances and plant hormones. Therefore, when artificially cultivating seedlings, first grading the seeds before sowing can improve the uniformity of the seedlings and reduce the labor cost of picking and grading seedlings.
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Keywords:
- Quercus variabilis /
- seed /
- soluble protein /
- abscisic acid /
- germination and emergence of seedlings
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图 1 不同规格种子贮藏物质含量
小写字母不同表示不同规格养分差异显著（P < 0.05）。Different small letters indicate significant difference of nutrients in different seed sizes (P < 0.05).
Figure 1. Content of storage substances in seeds of different specifications
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图 2 不同规格栓皮栎种子的 GA 3 （A）、IAA（B）和 ABA（C）含量
小写字母不同表示不同种子规格ABA含量差异显著（P<0.05）。Different small letters indicate significant difference in ABA content among different seed sizes (P<0.05).
Figure 2. Content of GA3 (A), IAA (B) and ABA (C) of Quercus variabilis seed with different sizes
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图 3 不同规格栓皮栎种子萌发动态图
不同小写字母代表不同规格种子萌发率之间差异显著（P<0.05）。Different lowercase letters represent significant difference in seed germination rates among varied specifications (P<0.05).
Figure 3. Dynamic diagram of seed germination of Q. variabilis of different specifications and sizes
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图 4 不同规格种子萌发率Probits统计量与萌发天数倒数的线性关系
Figure 4. Linear relationship between germination Probits statistics of seeds of different sizes and reciprocal days of geimination
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图 5 不同规格大小栓皮栎种子出苗动态图
不同小写字母代表不同规格种子出苗率之间差异显著（P<0.05）。Different lowercase letters represent significant difference in seedling emergence rates among seeds of different specifications (P<0.05).
Figure 5. Dynamic diagram of emergence of Q. variabilis seeds of different specifications and sizes
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图 6 不同规格种子出苗率Probits统计量与出苗天数倒数的线性关系
Figure 6. Linear relationship between emergence rate Probits statistics of seeds of different specifications and reciprocal days of emergence
表 1 栓皮栎不同规格种子单粒质量相关信息（n = 5 000）
Table 1 Information of the single-seed mass of different specifications of Quercus variabilis seeds (n = 5 000)
指标
Index小粒种子
Small seed （Ⅲ）中粒种子
Medium seed （Ⅱ）大粒种子
Large seed （Ⅰ）单粒质量范围 Single-seed mass range/g < 4.05 [4.05，5.46） ≥ 5.46 所占比例 Proportion/% 33.40 42.78 23.82 单粒质量最小值 Min. mass of single seed/g 0.23 4.05 5.46 单粒质量最大值 Max. mass of single seed/g 4.04 5.45 9.61 单粒质量均值 Mean mass of single seed/g 3.38 4.72 6.17 种子平均长度 Average length of seed/mm 21.100 0 ± 0.218 6c 22.300 0 ± 0.295 0b 23.500 0 ± 0.292 8a 种子平均宽度 Average width of seed/mm 16.800 0 ± 0.189 6c 18.600 0 ± 0.164 2b 20.700 0 ± 0.214 2a 种子平均长度/平均宽度 Average length/average width of seed 1.300 0 ± 0.016 3a 1.200 0 ± 0.021 1b 1.100 0 ± 0.017 5c 注：不同小写字母表示不同规格种子差异显著（P < 0.05）。Note: different small letters indicate significant difference between seeds of varied classes (P < 0.05). 
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表 2 影响栓皮栎种子萌发与出苗相关指标的主成分分析
Table 2 Principal component analysis of related indexes affecting seed germination and emergence of Q. quercus
主成分
Principal component可溶性糖
Soluble sugar淀粉
Starch可溶性蛋白
Soluble protein脱落酸
ABA赤霉素
GA3生长素
IAA赤霉素/脱落酸
GA3/ABA累计方差贡献率
Cumulative variance
contribution rate/%1 −0.007 0.183 0.239 −0.241 −0.087 0.190 0.203 66.344 2 −0.375 0.128 −0.201 0.153 0.411 0.109 0.073 100.000
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