Differences in Leaf Stomatal Traits across Plant Life Forms and Stomatal Distribution Types and Their Associations with Leaf Economic Traits

doi:10.7525/j.issn.1673-5102.2026.01.012

Abstract

Abstract:

To explore the differences in stomatal traits between different life forms（trees vs. shrubs） and stomatal distribution types（hypostomatous vs. amphistomatous）， and their relationships with leaf economic traits， this study was conducted from July to August 2023 using 22 common tree species on the campus of Northeast Forestry University. Through correlation analysis and standardized major axis analysis， the following findings were obtained：（1）Both life forms and stomatal distribution types significantly influenced stomatal traits. Trees exhibited significantly higher stomatal density（D_S）（P<0.05） but lower stomatal aperture（A_S） than shrubs. Hypostomatous leaves showed significantly higher D_S and stomatal relative area（A_SR） than amphistomatous leaves. （2）Relationships among stomatal traits varied by life form and leaf type. Negative correlations between D_S and stomatal size （S_S） or A_SR were statistically significant（P<0.05） only in trees and hypostomatous leaves， while the positive correlation between S_S and A_S reached significance（P<0.05） exclusively in hypostomatous leaves. （3）Stomatal traits were closely linked to the other leaf economic traits. Both D_S and A_SR were negatively correlated with specific leaf area （A_SL） and leaf nitrogen content（N_L）， but positively correlated with leaf dry matter content（C_LDM）. A_S showed a negative correlation with N_L. Trait coordination was more stable in trees and hypostomatous leaves， whereas shrubs and amphistomatous leaves exhibited more independent trait variations. The results of this research help to further understand the stomatal development strategies of different types of plants.

Key words: life form, amphistomatous leaves, stomatal traits, leaf economic traits

CLC Number:

S718.45

Yuhan LIU, Tingting LI, Liu YANG, Te LI, Gang LIU, Xiuwei WANG. Differences in Leaf Stomatal Traits across Plant Life Forms and Stomatal Distribution Types and Their Associations with Leaf Economic Traits[J]. Bulletin of Botanical Research, 2026, 46(1): 131-144.

Figures/Tables 10

Table 1

A summary of 22 common tree species investigated

气孔分布类型 Stomatal distribution type	科 Family	属 Genus	种 Species	生活型 Life form	胸径 DBH/cm	地径 DGL/cm
单面气孔型 Hypostomaty	无患子科Sapindaceae	槭属Acer	色木槭Acer pictum	乔木Tree	9.2~15.0
	无患子科Sapindaceae	槭属Acer	糖槭Acer saccharum	乔木Tree	14.5~19.0
	豆科Fabaceae	紫穗槐属Amorpha	紫穗槐Amorpha fruticosa	灌木Shrub		1.0~2.3
	桦木科Betulaceae	桦木属Betula	白桦Betula platyphylla	乔木Tree	20.2~25.6
	蔷薇科Rosaceae	山楂属Crataegus	山楂Crataegus pinnatifida	乔木Tree	9.6~18.5
	木樨科Oleaceae	梣属Fraxinus	水曲柳Fraxinus mandshurica	乔木Tree	32.3~37.2
	蔷薇科Rosaceae	苹果属Malus	海棠Malus spectabilis	乔木Tree	6.1~8.2
	木兰科Magnoliaceae	天女花属Oyama	天女木兰Oyama sieboldii	乔木Tree	3.3~5.7
	蔷薇科Rosaceae	稠李属Padus	紫叶稠李 Padus virginiana ‘Canada Red’	乔木Tree	9.1~18.3
	绣球科Hydrangeaceae	山梅花属 Philadelphus	东北山梅花 Philadelphus schrenkii	灌木Shrub		1.7~2.9
	蔷薇科Rosaceae	李属Prunus	稠李Prunus padus	乔木Tree	17.5~34.2
	蔷薇科Rosaceae	李属Prunus	榆叶梅Prunus triloba	灌木Shrub		1.3~2.9
	壳斗科 Fagaceae	栎属Quercus	蒙古栎Quercus mongolica	乔木Tree	12.3~15.4
	荚蒾科 Viburnaceae	接骨木属Sambucus	接骨木Sambucus williamsii	灌木Shrub		4.0~5.9
	蔷薇科 Rosaceae	花楸属Sorbus	花楸Sorbus pohuashanensis	乔木Tree	12.1~16.8
	锦葵科 Malvaceae	椴属Tilia	紫椴Tilia amurensis	乔木Tree	9.1~10.9
	锦葵科 Malvaceae	椴属Tilia	辽椴Tilia mandshurica	乔木Tree	12.3~22.1
双面气孔型 Amphistomaty	杨柳科Salicaceae	杨属Populus	钻天杨Populus nigra var. italica	乔木Tree	22.3~28.6
	杨柳科Salicaceae	杨属Populus	小叶杨Populus simonii	乔木Tree	50.5~55.5
	杨柳科Salicaceae	柳属Salix	旱柳Salix matsudana	乔木Tree	28.5~33.5
	木樨科Oleaceae	丁香属Syringa	暴马丁香Syringa reticulata	乔木Tree	3.6~9.0
	木樨科Oleaceae	丁香属Syringa	丁香Syringa oblata	灌木Shrub		3.2~6.7

Table 1

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统计指标 Statistical index	上表皮 Adaxial epidermis				下表皮 Abaxial epidermis
统计指标 Statistical index	气孔密度 D_S/（No.·mm^-2）	气孔大小 S_S/μm²	气孔相对面积A_SR/%	气孔开度 A_S/%	气孔密度 D_S/（No.·mm^-2）	气孔大小 S_S/μm²	气孔相对面积 A_SR/%	气孔开度 A_S/%
样本量 Sample size	24	24	24	24	109	109	109	109
平均值 Mean	38.06	342.87	1.22	48.53	276.75	347.95	7.41	46.14
最小值 Minimum	10.11	248.82	0.35	38.49	68.82	126.50	2.11	24.30
最大值 Maximum	97.69	605.41	2.47	55.85	1 057.21	1 386.50	22.49	60.21
标准误差 Standard error	4.67	18.56	0.12	0.81	21.69	21.47	0.40	0.72
偏度 Skewness	1.18	1.27	0.61	-0.74	1.70	2.55	1.50	-0.32
峰度 Kurtosis	0.63	1.52	-0.77	0.76	2.07	8.13	2.39	-0.26
变异系数 Coefficient of variation	0.60	0.27	0.50	0.08	0.82	0.64	0.57	0.16

性状 Trait	效应 Effect	自由度 d_f	F	P
气孔密度 D_S	生活型 Life form	1	7.91	**
	气孔分布类型 Stomatal distribution type	1	2.31	0.13
	气孔分布类型×生活型 Stomatal distribution type×Life form	1	5.44	*
气孔大小 S_S	生活型 Life form	1	3.40	0.07
	气孔分布类型 Stomatal distribution type	1	0.40	0.53
	气孔分布类型×生活型 Stomatal distribution type×Life form	1	0.01	0.93
气孔相对面积 A_SR	生活型 Life form	1	3.50	0.07
	气孔分布类型 Stomatal distribution type	1	7.50	**
	气孔分布类型×生活型 Stomatal distribution type×Life form	1	10.70	**
气孔开度 A_S	生活型 Life form	1	5.86	*
	气孔分布类型 Stomatal distribution type	1	3.95	*
	气孔分布类型×生活型 Stomatal distribution type×Life form	1	0.23	0.63

特征 Trait	叶资源经济性状 Leaf economic traits						气孔特征 Stomatal traits
特征 Trait	比叶面积 A_SL	叶面积 A_L	叶片厚度 T_L	叶绿素含量 Chl	叶片氮含量 N_L	干物质含量 C_LDM	气孔密度 D_S	气孔大小 S_S	气孔相对面积 A_SR	气孔开度 A_S
气孔密度D_S	-0.31^**	0.03	-0.04	-0.03	-0.19^*	0.51^**	1
气孔大小S_S	-0.03	0.73	-0.01	-0.14	-0.18	-0.13	-0.44^**	1
气孔相对面积A_SR	-0.37^**	0.13	0.05	-0.15	-0.36^**	0.42^**	0.77^**	0.12	1
气孔开度A_S	-0.18	-0.04	0.07	0.01	-0.21^*	0.07	-0.20^*	0.40^**	0.10	1

性状 Trait	生活型 Life form				气孔分布类型 Stomatal distribution type
	乔木 Tree		灌木 Shrub		单面气孔型 Hypostomaty		双面气孔型 Amphistomaty
	R²	P	R²	P	R²	P	R²	P
D_S-S_S	0.50	**	0.27	0.37	0.53	***	0.16	0.50
D_S-A_SR	0.53	***	0	0.95	0.45	**	0.46	0.21
A_S-S_S	0.16	0.11	0.36	0.28	0.25	*	0.09	0.62