基于4-PHF（the four-parameter Hill function)法比较6种四照花种子的休眠特性

熊星; 鲁强; 王昊伟; 杨玲; 洑香香

doi:10.19303/j.issn.1008-0384.2022.004.008

基于4-PHF（the four-parameter Hill function)法比较6种四照花种子的休眠特性

doi: 10.19303/j.issn.1008-0384.2022.004.008

南京林业大学南方现代林业协同创新中心/南京林业大学林学院，江苏南京 210037

基金项目: 江苏省林业科技创新与推广项目（LYKJ[2018]06）

详细信息

作者简介:
熊星（1996−），女，硕士研究生，主要从事森林生态学研究（E-mail：1366309615@qq.com）

通讯作者:
洑香香（1969−），女，教授，博士生导师，从事森林培育研究（E-mail：xxfu@njfu.edu.cn）

中图分类号: S 68
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出版历程
- 收稿日期: 2021-11-03
- 修回日期: 2022-04-11
- 网络出版日期: 2022-04-24
- 刊出日期: 2022-04-28

4-PHF-based Analysis on Seed Dormancy of 6 Cornus Species

Co-Innovation Center for Sustainable Forestry in Southern China, Nanjing Forestry University/College of Forestry, Nanjing Forestry University, Nanjing, Jiangsu　210037, China

摘要

摘要: 目的四照花属（Cornus）树种具有很高的园林观赏价值，有较高开发利用前景。本研究利用4-PHF（the four-parameter Hill function）法比较6种四照花种子的休眠特性，为四照花种质资源保存、种子贮藏及开发利用等提供理论依据。方法以6种四照花，包括大花四照花（CF）、中国四照花（CKC）、香港四照花（CH）、东京四照花（CHT）、秀丽四照花（CHE）和尖叶四照花（CE）种子为材料，用500 mg·L^-1赤霉素（GA₃）溶液浸泡3 d后进行低温层积处理，并设7个层积时间梯度（0、10、20、30、40、50、60 d）；采用4-PHF法拟合6种四照花种子发芽曲线，计算发芽参数包括起始发芽时间（lag）和发芽高峰时间（TMGR）及休眠指数（DI）。结果层积有效地促进了四照花种子休眠的解除，但每个种达到最大发芽率所需层积时间存在差异：CKC仅需层积30 d发芽率可达78%，CH和CF层积60 d后发芽率分别达71%和89%，CHE、CHT和CE层积50 d后发芽率分别达63%、69%和54%。用4-PHF方程拟合的发芽曲线表明：随着休眠的解除，发芽参数lag和TMGR值持续下降，DI则呈现增加的趋势。结论结合各个发芽参数得出落叶类四照花CF、CKC种子休眠程度较浅，而常绿种CH、CHT、CHE和CE种子休眠较深，由DI值得出本试验处理未能充分解除常绿类四照花种子休眠，4- PHF方程进行种子萌发曲线拟合及萌发参数lag和TMGR可从生物学角度诠释层积处理对6种四照花种子的萌发差异，但DI值不宜用于比较种间休眠程度的差异。
- 四照花属 /
- 起始发芽时间（lag） /
- 发芽高峰时间（TMGR） /
- 休眠指数（DI）
Abstract: Objectives Seed dormancy of 6 species of ornamental Cornus was studied using the 4-parameter hill function (4-PHF) method to aid the resource conservation, seed storage, plant development, and landscape planning. Method Seeds of C. florida (CF), C. kousa subsp. chinensis (CKC), C. hongkongensis (CH), C. hongdongensis subsp. tonkinensis (CHT), C. hongdongensis subsp. elegans (CHE), and C. elliptica (CE) were soaked in a 500 mg·L⁻¹ GA₃ solution for 3 d. Then, at durations of 0, 10, 20, 30, 40, 50, and 60 d, the seeds were stratified at low temperature. The 4-PHF method was used for curve-fitting to describe germinating patterns, as well as calculating estimates on the time at germination onset (lag), time at maximum germination rate (TMGR), and dormancy index (DI) for the 6 species. Result The stratification effectively facilitated the break of seed dormancy. However, the holding time required for maximal germination varied among the 6 species. For CKC, 78% germination was achieved in 30 d; for CH and CF, it was 71% and 89%, respectively, in 60 d; and for CHE, CHT, and CE, it was 63%, 69%, and 54%, respectively, in 50 d. The germination curves showed steady declines after break of dormancy on lag and TMGR but increase on DI. Conclusion As indicated by the germination parameters, it appeared that the evergreen variety of dogwoods went into a deeper dormancy than the deciduous ones. The DI showed the current treatment could not fully break the seed dormancy on the evergreen dogwoods. The germination curve-fitting and the 4-PHF-estimated lag and TMGR could explain the biological differences on the seed germination among the 6 dogwood species by the stratification treatments. On the other hand, DI could not be used to compare the degree of dormancy.
- Cornus /
- time at germination onset /
- time at maximum germination rate /
- dormancy index

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图 1 层积时间对各种四照花种子绝对发芽率的影响

注：不同大写字母表示同种四照花种子经不同层积时间处理后的发芽率差异显著性(P<0.05)；小写字母表示不同种四照花种子相同层积处理时间的种子发芽率差异显著性(P<0.05)。

Figure 1. Effect of stratification duration on absolute seed germination percentage of various dogwoods

Note: Data with different capital letters indicate significant difference on germination rate of same kind of dogwoods under varied stratification duration (P＜0.05); those with different lowercase letters indicate significant difference on germination rate of different species of dogwoods under same stratification treatment (P＜0.05).

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图 2 6种四照花种子不同层积时间下累积发芽率的4-PHF拟合曲线

注：符号代表随时间推移的累积发芽率，实线代表拟合曲线。

Figure 2. 4-PHF fitting curves for cumulative germination rates of dogwood seeds after stratification

Note: Symbols represent cumulative germination over time; solid lines are fitted curves.

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图 3 6种四照花种子不同层积时间下的瞬时萌发率

注：曲线上方数字为对应发芽曲线的TMGR值。

Figure 3. Instant germination rate of dogwood seeds after stratification

Note: Data above curve are corresponding TMGR.

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图 4 6种四照花种子的休眠指数（DI）比较

注：y1(上曲线)分别为大花四照花和香港四照花层积60 d，秀丽四照花、东京四照花和尖叶四照花层积50 d，中国四照花层积30 d的萌发曲线；y2(下曲线)为未层积处理的萌发曲线。阴影面积代表其休眠指数DI。

Figure 4. DIs for seeds of 6 dogwood species

Note: DI: shaded area between Y₁ (upper curves) of germination curves of CF stratified for 60 d, CH for 60 d, CHE for 50 d, CHT for 50 d, CE for 50 d, and CKC for 30 d and Y₂ (lower curve) of those of the unstratified counterparts.

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表 1 6种四照花种子采集信息

Table 1. Information on seed collection

树种名称 Specific name	代号 Code name	千粒重 Thousand seed weight/g	采种位置 Seed position	经纬度 Longitude and latitude	海拔高度 Altitude/m
中国四照花 C. kousa subsp. chinensis	CKC	44.998	甘肃省天水市麦积区马跑泉镇崖湾村 Maiji District, Tianshui, Gansu Province	E 105°54′32"，N 34°30′33"	1170
秀丽四照花 C. hongdongensis subsp. elegans	CHE	61.000	浙江龙泉林科院 Longquan Forestry Research Institute	E 119°6′16"，N 28°2′7"	424
东京四照花 C. hongdongensis subsp. tonkinensis	CHT	83.045	南京市溧水区四照花园 Lishui District, Nanjing, Jiangsu Province	E 119°2′4"，N 31°35′14"	81
香港四照花 C. hongkongensis	CH	60.000	广东省韶关市乳源瑶族自治县洛阳镇 Ruyuan Yao Autonomous County, Shaoguan, Guangdong Province	E 113°2′38"，N 24°39′47"	700
尖叶四照花 C. elliptica	CE	70.000	广东省韶关市乐昌市廊田镇白山村 Lechang City, Shaoguan, Guangdong Province	E 113°28′8"，N 25°7′29"	160
大花四照花 C. florida	CF	95.999	美国路易斯安那州 State of Louisiana	W 90° 03′00″， N 29° 58′00″	30

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表 2 不同层积时间下6种四照花种子休眠解除的四参数希尔方程（4-PHF）的拟合参数

Table 2. 4-PHF fitting parameters for dormancy release of dog-wood seeds after stratification

代号 Code	参数 Parameter	层积时间 Stratification duration/d
代号 Code	参数 Parameter	0	10	20	30	40	50	60
CF	a	49.02	70.02	76.12	78.57	79.17	81.63	115.80
	b	3.93	4.29	4.73	4.96	4.97	3.24	2.16
	c	17.94	15.46	11.64	7.55	7.52	7.00	3.24
	y₀	0.53	−0.96	−1.15	−1.33	−1.33	0.66	−25.41
CKC	a	19.07	52.54	66.54	76.97	46.17	54.59	45.52
	b	5.52	6.20	4.93	6.40	2.73	1.87	3.50
	c	13.83	11.36	10.44	12.21	11.13	11.17	6.38
	y₀	−0.46	−0.52	−0.82	0.96	−1.52	−3.90	3.34
CHE	a	−	−	−	6.48	1611.00	65.25	57.45
	b	−	−	−	5.92	2.92	2.97	2.09
	c	−	−	−	12.59	12.90	7.69	5.54
	y₀	−	−	−	1.22	2.29	−0.17	−1.54
CHT	a	5.20	8.18	11.48	23.45	57.99	72.35	62.49
	b	12.74	7.49	5.74	3.04	1.98	2.66	2.67
	c	18.63	15.93	17.71	15.47	5.61	4.59	11.69
	y₀	−0.14	−0.03	−0.13	−0.70	−17.10	−2.67	2.34
CH	a	11.59	31.31	44.43	55.79	65.51	73.93	75.96
	b	6.71	4.78	4.08	3.42	2.82	2.11	2.94
	c	19.05	22.65	19.55	22.18	15.49	12.28	13.32
	y₀	−0.38	−2.83	−1.06	0.06	−1.76	−2.28	0.89
CE	a	14.08	25.53	40.82	41.19	50.22	56.65	53.16
	b	7.70	3.25	2.28	2.16	2.22	1.84	1.77
	c	17.75	15.01	11.40	11.20	8.48	4.41	5.92
	y₀	0.23	−2.13	−4.53	−2.88	−1.40	−2.21	0.20

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表 3 不同层积时间下6种四照花种子的初始萌发时间（lag）的实际值与理论值

Table 3. Actual and theoretical lag of dogwood seeds after stratification

层积时间 Stratification duration/d	CF	CKC	CHE	CHT	CH	CE
0	*/5	7.07/8	0	14.05/16	11.49/13	*/11
10	5.71/7	5.41/7	0	7.54/10	13.97/15	7.17/9
20	4.82/6	4.30/6	0	8.15/10	7.88/9	4.57/6
30	3.33/5	*/6	*/6	4.94/7	*/7	3.37/4
40	3.31/5	3.22/5	*/4	5.54/5	4.34/5	1.71/3
50	*/1	2.83/5	1.03/2	1.34/2	2.41/3	0.77/1
60	1.80/1	*/2	1.00/2	1.29/2	*/3	*/1
注： “*”代表求解无实根，即萌发曲线与时间轴（x轴）无交点;“/”前面表示理论的初始萌发时间，后面表示实际的初始萌发时间。
Note: ″*″ represents solution without real roots, i.e., germination curve does not intersect time (x-axis). Data in front of ″/″ are theoretical initial germination duration, and those behind “/”, actual time.

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