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紫色土丘陵区是典型亚热带季风气候的生态脆弱区[1],受城市化建设规模加速扩大的影响,开山修路、削山平地等建设措施改变了原有高低起伏的地形地貌,形成了诸多裸露边坡[2],破坏了原有地带性植被。为稳定边坡、恢复植被和增加景观,常常以建植草坪的方式进行边坡治理。但建成的边坡草坪因地形地势无法进行精细的养护,又受气候特点、土层厚度及草坪草生态学特征的限制,2~3年后退化殆尽[3],逐步演替成耐受性强的荩草(Arthraxon hispidus)、五节芒(Miscanthus floridulus)等草本物种为优势种的次生群落[4]。构树(Broussonetia papyrifera)是边坡草坪中次生演替的木本植物,木本植物因其根深等特点比草本植物具有更强的固坡护土能力,对生态系统产生更为深远的影响。故开展边坡草坪中的次生构树对边坡生境适应性及生长策略研究,可为该区裸露边坡植被恢复时植物选择与配置提供借鉴。
构树为桑科(Moraceae)构树属(Broussonetia)雌雄异株植物,具材用、饲用、药用、食用等诸多价值有待开发利用。作为木本饲料、受损生境修复和生态治理乡土树种,近年来诸多学者对其形态特征、群落分布特点、化学成分、饲用及药用价值[5]及高温干旱、盐碱及重金属等胁迫下的适应性和抗性进行了大量研究[6-9]。但常忽略其性别分化的雌雄异株特性,仅对抗旱性[10]、叶片构造[11]和同工酶[12]的性别差异进行了比较。受遗传特性、生理特化、环境因子以及生殖成本共同作用[13],使雌雄异株植物个体形态塑性、生理响应、生长和繁殖策略及生活史等方面存在差异[14-15],导致雌雄个体对生境胁迫的响应速度、调节能力和适应策略具有性别差异[16],表现为适应性、抗逆性及生理反应的性别差异。从而影响个体生长、种群特征和群落结构,甚至可能影响植物物种的延续、生态系统的退化速度及演替方向。
本研究以人工建植6年后的边坡草坪为样地,划定坡上、坡中、坡下及坡底4个坡位,以次生演替的构树幼树种群为材料,通过测定次生构树种群的年龄结构、性比组成及分布系数等指标,分析坡位对构树种群特征及个体生长的影响,拟回答(1)坡位是否影响次生构树种群数、种群大小、年龄结构、性比、分布系数等种群特征参数?(2)雌雄个体是否对坡位生境采取不同的生长策略而体现出抗性差异?(3)构树是否适合在裸露边坡植被恢复中推广应用?以期为裸露边坡植被恢复、构树品种选育提供依据,为雌雄异株植物个体发育、种群形成研究及开发和利用途径提供参考资料。
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4个坡位的坡度存在极显著差异(P<0.01)(见表1),随坡位降低,土层厚度增加、土壤容重降低、有机质含量和含水量增加(P<0.05),使4个坡位的土层厚度、土壤容重、有机质含量及土壤含水量存在极显著差异(P<0.01)。坡下的土层厚度显著高于其他坡位,其有机质含量与坡底差异较小,但显著大于坡上和坡中。因坡度影响水土的截留能力,分析F值可知,坡位对土壤性状影响大小为:土壤含水量>土层厚度>土壤容重>有机质含量。土壤含水量为差异最大的生态因子,使不同坡位上构树幼苗种群处于不同干旱胁迫生境中。
表 1 种群坡度及土壤性状
Table 1. Slope position and soil properties
坡位
Slope position坡度
Slope /˚土层厚度
Soil thickness/cm土壤容重
Soil bulk density/(g·cm–3)土壤含水量
Soil water content/%有机质含量
Organic matter content /%坡上Uphill 35.00±3.64a 7.21±1.33d 1.85±0.25a 21.14±1.24c 1.02±0.24c 坡中Middle 25.00±2.51b 13.65±2.67c 1.68±0.14b 32.53±1.98bc 1.83±0.42b 坡下Downhill 15.00±1.32c 34.67±10.49a 1.47±0.03c 39.26±3.92b 2.52±0.32a 坡底Flat 5.00±1.22d 22.62±4.35b 1.46±0.42c 45.66±1.37a 2.63±0.17a F 32.337 24.631 8.623 28.274 7.236 P <0.001 <0.001 <0.001 <0.001 <0.001 注:同列不同小写字母表示坡位间的差异显著(P<0.05)。表中数值为平均值±标准差。下表同
Note: Different lowercase letters in the same column indicate significant differences in slope position (P<0.05). The values in the table are average±standard deviation. The same as below. -
坡位对构树幼树种群面积有显著影响(P<0.05),对种群中一年生苗、幼树和成年树植株比例有极显著影响(P<0.01)(见表2)。构树的种群面积、种群密度随坡位从高到低逐步增加,坡底种群面积和种群密度最大。坡底种群面积为坡上、坡中、坡下的8.24倍、5.06和1.48倍,密度分别为坡上、坡中和坡下的3.57倍、1.89倍和1.28倍。由F值可见,坡位对种群密度影响大于种群面积。
表 2 坡位对种群特征的影响
Table 2. Effects of slope position on population area, population density and age structure
坡位
Slope position种群面积
Population area/m2种群密度
Population density/
(plant·m–2)年龄结构 Age structure/% 一年生苗
One-year seedling幼苗
Seedling幼树
Youth tree成年树
Mature tree坡上Uphill 4.21±0.37d 2.33±0.33d 6.52±3.14d 25.52±7.03b 34.61±9.32a 33.35±6.32b 坡中Middle 6.86±1.68c 4.35±1.33c 8.37±2.26c 26.35±6.35b 26.89±7.87b 38.39±9.17a 坡下Downhill 23.38±9.67b 6.42±1.67b 13.64±3.41b 32.66±2.68a 26.32±6.32b 27.38±6.28c 坡底Flat 34.68±11.27a 8.24±1.33a 40.38±6.52a 25.27±4.05b 13.62±4.34c 20.73±4.08d F 26.541 42.612 48.177 4.274 16.374 8.315 P <0.001 <0.001 <0.001 0.031 <0.001 <0.001 随坡位降低,一年生苗的比例显著增加,幼树和成年树的比例显著降低(P<0.05)。坡下幼苗比例显著高于其他坡位,而坡上、坡中和坡底间差异较小。坡底一年生苗比例最大,坡下幼苗比例最大。坡位对年龄结构有显著影响,坡上和坡中种群表现出衰退型特点,坡下种群相对稳定,坡底种群为增长型。坡位对年龄结构影响大小为一年生苗>幼树>成年树>幼苗。
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坡位对一年生苗、幼苗、幼树及成年树植株的株高均有极显著影响(P<0.01)(见表3),随坡位降低各龄构树的株高均逐步增加。坡位对株高影响随植株年龄增大而增大,坡底一年生苗、幼苗、幼树及成年树构树株高分别是坡上、中、下的2.17倍、1.87倍、1.25倍,2.68倍、1.85倍、1.34倍,1.86倍、1.68倍、1.08倍和2.64倍、2.14倍、1.09倍。可见坡位对一年生苗株高影响较小,对幼苗和成年树构树株高影响较大。
表 3 坡位对个体大小的影响
Table 3. Effect of slope position on individual size
坡位
Slope position株高 Plant height/cm 地径 Stem diameter /cm 一年生苗
One-year
seedling幼苗
Seedling幼树
Youth tree成年树
Mature tree一年生苗
One-year
seedling幼苗
Seedling幼树
Youth tree成年树
Mature tree坡上Uphill 9.71±1.04d 15.33±3.12d 70.75±10.11d 81.68±20.31d 0.28±0.0b 0.66±0.09d 3.21±0.26c 5.24±0.12c 坡中Middle 11.27±1.88c 22.21±6.21c 78.46±9.63c 100.37±26.41c 0.30±0.0b 1.07±0.47c 3.68±0.31b 6.12±1.11b 坡下Downhill 16.86±2.11b 30.62±9.24b 121.32±24.18b 198.42±41.62b 0.29±0.0b 2.09±1.02a 4.42±0.21a 6.96±1.40a 坡底Flat 21.12±6.12a 41.09±8.17a 132.04±35.07a 215.24±37.85a 0.45±0.16a 1.63±1.21b 3.57±0.49b 5.97±0.51b F 37.285 43.527 57.818 81.124 4.246 15.857 4.817 4.121 P <0.001 <0.001 <0.001 <0.001 0.039 <0.001 0.022 0.044 坡位对一年生苗、幼苗、幼树及成年树的地径均有显著影响(P<0.05),对幼苗地径的影响最大,对幼树影响次之,对成年树影响最小(表3)。坡上各龄植株的地径显著低于其他坡位,坡下幼苗、幼树及成年树的地径均为最大。坡底一年生苗地径显著高于其他地形,而坡上、中、下间无显著差异。坡上与坡底植株地径随树龄增加差异逐渐缩小。坡位对地径影响大小为幼苗>幼树>一年生苗>成年树。比较F值可知,坡位对株高的影响大于地径。
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坡位对幼树和成年树的雌雄性比有极显著影响(P<0.01),性比随坡位降低显著增加(P<0.05)(表4)。坡上和坡中的幼树和成年树的雌雄性比均小于1,雌株数显著小于雄株数。坡上幼树和成年树的性比差异较小,坡中幼树的性比显著大于成年树,坡下和坡底幼树和成年树的雌雄性比均大于1。不同坡位幼树、成年树雌雄性比的差异表明,高坡位雄株比例大,其抗旱性强于雌株,而低坡位雌株对空间等其他资源竞争力高于雌株。
表 4 坡位对种群性比与分布系数的影响
Table 4. Effects of slope position on sex ratio and distribution coefficient of population
坡位
Slope position性比 Sex ratio(♀:♂) 分布系数 Distribution coefficient 幼树
Youth tree成年树
Mature tree一年生苗
One-year seedling幼苗
Seedling幼树 Youth tree 成年树 Mature tree ♀ ♂ ♀ ♂ 坡上 Uphill 0.25±0.04d 0.27±0.09d 1.63±1.42c 1.23±0.09c 1.53±0.04b 1.51±0.04a 1.04±0.06a 1.04±0.06a 坡中 Middle 0.46±0.12c 0.35±0.13c 4.62±1.25b 2.08±0.47b 1.56±0.05b 1.42±0.05a 1.02±0.04a 1.02±0.04a 坡下 Downhill 1.32±0.07b 3.42±1.33b 5.77±1.05a 3.19±1.02a 2.42±0.18a 1.12±0.18b 0.55±0.31b 1.05±0.31a 坡底 Flat 2.78±0.82a 5.24±1.67a 6.15±1.44a 3.24±1.21a 2.57±0.23a 1.08±0.23b 0.27±0.24b 0.97±0.24a F 27.624 32.416 16.724 5.238 4.243 3.834 5.121 2.121 P <0.001 <0.001 <0.001 0.017 0.042 0.046 0.026 0.768 -
坡位对一年生苗、幼苗和幼树的分布系数有显著影响(P<0.05)(见表4),对成年雌株Cx有显著影响,对雄株无影响。一年生苗、幼苗和幼树的Cx随年龄增加逐渐降低。4个坡位上,随树龄增加Cx均表现为下降,表明种群发展过程中,每个阶段都有各龄植株死亡。
一年生苗和幼苗树Cx均大于1,且随坡位降低而增加,坡位越低一年生苗和幼苗树越呈集群分布。幼树和成年树Cx表现出显著的性别差异,幼树雌株Cx随坡位降低而增加,雄株Cx则下降;成年雌株随坡位降低Cx降低,而雄株Cx则无变化。随树龄增加分布系数逐步下降,成年树时分布系数近于1或0,由集群分布逐步变为随机分布或均匀分布。成年树雌株随坡位降低呈均匀分布,坡位对成年树雄株Cx影响较小且接近于随机分布。坡位对分布系数影响大小为一年生苗>幼苗>成年树雌株>幼树雌株>幼树雄株>成年树雄株。
Effects of Slope Position on Population Characteristics and Growth Strategies of Dioecious Broussonetia papyrifera in Degraded Slope Lawn
More Information-
摘要: 为了研究雌雄个体对自然坡位适应性的性别差异,以紫色土丘陵区人工建植6年后的3个坡向的边坡草坪为对象,通过测定坡上、坡中、坡下及坡底4个坡位上次生演替构树幼树种群的种群面积、种群密度、年龄结构、雌雄性比、分布系数等特征参数及雌雄个体株高和地径,研究坡位对次生构树种群特征和雌雄个体生长的影响。结果表明:(1)4个坡位的坡度和土壤理化性状有显著差异(P<0.05),构树均可入侵形成次生种群,随坡位降低种群面积、种群密度成倍增加(P<0.05);(2)坡位显著影响种群年龄结构,随坡位降低一年生苗树比例增加,幼树和成年树比例降低(P<0.05),坡上和坡中年龄结构为衰退型,下坡为稳定型,坡底为增长型;(3)坡位显著影响幼树和成年树的性比,坡上和坡中偏向雄株,下坡和坡底偏向雌株,坡位对成年树性比影响大于幼树;(4)坡位显著影响各龄构树的空间分布格局,坡位越低一年生苗和幼苗树越呈集群分布,随树龄增加分布系数均为下降,雌株趋向均匀分布,雄株趋向随机分布;(5)坡位对各龄构树的株高影响大于地径,株高随坡位降低而增加,地径随土层厚度增加而增加。综上,边坡草坪演替成构树+草本的混生群落,雄株对资源匮乏的耐受力强于雌株,雌株对资源的竞争力强于雄株,表现为与坡位的坡向和土壤理化性质相匹配的次生种群特征和个体生长策略,适合在裸露边坡植被恢复中推广与应用。Abstract: In order to study the gender differences of male and female individuals' adaptability to natural slope position, the secondary Broussonetia papyrifera population in the degraded slope lawn of purple hilly area was used as study materials. The slope was divided into four slope positions as uphill, middle, downhill and flat, and population characteristics such as population area, population density, age structure, male-female ratio, distribution coefficient, and other characteristic parameters, as well as the height and ground diameter of male and female individuals were determined. The results showed that: (1) There were significant differences in slope size and soil physical and chemical properties among the 4 slope positions (P<0.05). B. papyrifera could invade each position and form secondary population, but the population area and density increased exponentially with the decrease of slope position (P<0.05). (2) The slope position significantly influenced the age structure of the population. With the decrease of the slope position, the proportion of one-year seedlings increased, while the proportion of young trees and adult trees decreased (P<0.05). The age structure of B. papyrifera population on uphill and middle slope was declining type, the downhill was stable and the flat was increasing type. (3) The slope position significantly affected the sex ratio of young trees and adult trees, with male plants on the uphill and middle slopes, and female plants on the downhill and flat slopes. (4) The slope position significantly affected the age spatial distribution pattern of B. papyrifera. The lower the slope position, the more annual seedlings and seedling trees were clustered. With the increase of tree age, the distribution coefficient decreased, and the females tended to be evenly distributed, while the males tended to be randomly distributed. (5) The slope position had more influence on the height of different trees than the ground diameter. The plant height increased with the decrease of slope position, while the ground diameter increased with the increase of soil thickness. To sum up, B. papyrifera invaded the degraded lawn and formed a mixed community of B. papyrifera and herbs. The males' tolerance to the lack of resources was better than females, and the females' competitiveness to resources was stronger than males. The secondary population characteristics and individual growth strategy matched with the slope position and soil physical and chemical properties, so B. papyrifera was suitable for popularization and application in vegetation restoration of bare slope.
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表 1 种群坡度及土壤性状
Tab. 1 Slope position and soil properties
坡位
Slope position坡度
Slope /˚土层厚度
Soil thickness/cm土壤容重
Soil bulk density/(g·cm–3)土壤含水量
Soil water content/%有机质含量
Organic matter content /%坡上Uphill 35.00±3.64a 7.21±1.33d 1.85±0.25a 21.14±1.24c 1.02±0.24c 坡中Middle 25.00±2.51b 13.65±2.67c 1.68±0.14b 32.53±1.98bc 1.83±0.42b 坡下Downhill 15.00±1.32c 34.67±10.49a 1.47±0.03c 39.26±3.92b 2.52±0.32a 坡底Flat 5.00±1.22d 22.62±4.35b 1.46±0.42c 45.66±1.37a 2.63±0.17a F 32.337 24.631 8.623 28.274 7.236 P <0.001 <0.001 <0.001 <0.001 <0.001 注:同列不同小写字母表示坡位间的差异显著(P<0.05)。表中数值为平均值±标准差。下表同
Note: Different lowercase letters in the same column indicate significant differences in slope position (P<0.05). The values in the table are average±standard deviation. The same as below.表 2 坡位对种群特征的影响
Tab. 2 Effects of slope position on population area, population density and age structure
坡位
Slope position种群面积
Population area/m2种群密度
Population density/
(plant·m–2)年龄结构 Age structure/% 一年生苗
One-year seedling幼苗
Seedling幼树
Youth tree成年树
Mature tree坡上Uphill 4.21±0.37d 2.33±0.33d 6.52±3.14d 25.52±7.03b 34.61±9.32a 33.35±6.32b 坡中Middle 6.86±1.68c 4.35±1.33c 8.37±2.26c 26.35±6.35b 26.89±7.87b 38.39±9.17a 坡下Downhill 23.38±9.67b 6.42±1.67b 13.64±3.41b 32.66±2.68a 26.32±6.32b 27.38±6.28c 坡底Flat 34.68±11.27a 8.24±1.33a 40.38±6.52a 25.27±4.05b 13.62±4.34c 20.73±4.08d F 26.541 42.612 48.177 4.274 16.374 8.315 P <0.001 <0.001 <0.001 0.031 <0.001 <0.001 表 3 坡位对个体大小的影响
Tab. 3 Effect of slope position on individual size
坡位
Slope position株高 Plant height/cm 地径 Stem diameter /cm 一年生苗
One-year
seedling幼苗
Seedling幼树
Youth tree成年树
Mature tree一年生苗
One-year
seedling幼苗
Seedling幼树
Youth tree成年树
Mature tree坡上Uphill 9.71±1.04d 15.33±3.12d 70.75±10.11d 81.68±20.31d 0.28±0.0b 0.66±0.09d 3.21±0.26c 5.24±0.12c 坡中Middle 11.27±1.88c 22.21±6.21c 78.46±9.63c 100.37±26.41c 0.30±0.0b 1.07±0.47c 3.68±0.31b 6.12±1.11b 坡下Downhill 16.86±2.11b 30.62±9.24b 121.32±24.18b 198.42±41.62b 0.29±0.0b 2.09±1.02a 4.42±0.21a 6.96±1.40a 坡底Flat 21.12±6.12a 41.09±8.17a 132.04±35.07a 215.24±37.85a 0.45±0.16a 1.63±1.21b 3.57±0.49b 5.97±0.51b F 37.285 43.527 57.818 81.124 4.246 15.857 4.817 4.121 P <0.001 <0.001 <0.001 <0.001 0.039 <0.001 0.022 0.044 表 4 坡位对种群性比与分布系数的影响
Tab. 4 Effects of slope position on sex ratio and distribution coefficient of population
坡位
Slope position性比 Sex ratio(♀:♂) 分布系数 Distribution coefficient 幼树
Youth tree成年树
Mature tree一年生苗
One-year seedling幼苗
Seedling幼树 Youth tree 成年树 Mature tree ♀ ♂ ♀ ♂ 坡上 Uphill 0.25±0.04d 0.27±0.09d 1.63±1.42c 1.23±0.09c 1.53±0.04b 1.51±0.04a 1.04±0.06a 1.04±0.06a 坡中 Middle 0.46±0.12c 0.35±0.13c 4.62±1.25b 2.08±0.47b 1.56±0.05b 1.42±0.05a 1.02±0.04a 1.02±0.04a 坡下 Downhill 1.32±0.07b 3.42±1.33b 5.77±1.05a 3.19±1.02a 2.42±0.18a 1.12±0.18b 0.55±0.31b 1.05±0.31a 坡底 Flat 2.78±0.82a 5.24±1.67a 6.15±1.44a 3.24±1.21a 2.57±0.23a 1.08±0.23b 0.27±0.24b 0.97±0.24a F 27.624 32.416 16.724 5.238 4.243 3.834 5.121 2.121 P <0.001 <0.001 <0.001 0.017 0.042 0.046 0.026 0.768 -
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