浙江人工红树林对关键环境因子的生态响应研究
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摘要
造林试验表明,温度、盐度和滩位是浙江人工红树林造林成功与否的主要限制因子。开展红树植物对温度、盐度和滩位的生态响应研究,将为人工红树林树种选择适宜的立地条件提供理论依据与技术支撑。在我国红树植物引种的北缘地区——浙江,开展人工红树林对关键环境因子的生态响应研究,具有重要科学价值。本文以我国亚热带地区优势红树植物秋茄(Kandelia obovata)及近几年浙江引种的桐花树(Aegiceras corniculatum)、无瓣海桑(Sonneratia apetala)等10种红树植物为材料,通过野外造林试验和温室模拟实验,采用野外调查监测和实验室分析、定性和定量相结合、单项指标和综合分析的方法,比较受试植物的形态指标、滩位适应度、寒害指数、生理指标、光合指标的变化,分析其在浙南海岸对环境因子的适应能力,系统研究了温度、盐度、滩位等关键环境因子对浙江人工红树林发展的影响,以及不同滩位生境中不同林龄秋茄种群结构和生物量模型。研究结果明确了全球气候变化背景下适宜浙江引种的红树植物种类和宜林生境。
(1)在全球气候变化背景下,对浙江红树林分布状况进行了预测。若年均温升温1.8℃,我国红树植物分布北界由福建福鼎向北推延至浙江舟山,分布到浙江的红树植物将达到3种,为秋茄、白骨壤(Avicennia marina)、桐花树。若升温4.0℃℃,我国红树植物分布北界将向北推延至南京,分布到浙江的红树植物将达到12种,为秋茄、白骨壤、桐花树、海漆(Excoecaria agallocha)、无瓣海桑等。该工作对浙江人工红树林造林树种的引选具有理论指导意义。
(2)浙江人工红树林造林树种对温度的响应。低温尤其是极端低温严重影响红树植物的生长。随着温度的降低,受试植物叶片相对电导率(RE)、脯氨酸含量(PRO)、超氧化物歧化酶活性(SOD)、丙二醛含量(MDA)逐渐增大。越冬后,拉关木(Laguncularia racemosa)和白骨壤全部死亡(寒害指数,Cii>3),无瓣海桑部分死亡(Cii=2.76),桐花树和秋茄的寒害指数与乡土植物无柄小叶榕(Ficuas concinna var. subsessilis)相似(1.08-1.14),无冻害现象。-4℃C极端低温过后,保存率依次为秋茄(77%)>桐花树(73%)>海漆(70%)>无瓣海桑(25%),拉关木、木榄(Bruguiera gymnorhiza)、尖瓣海莲(Bruguiera sexangula var. rhynchopetala)、红海榄(Rhizophora stylosa)、老鼠簕(Acanthus ilicifolius)、白骨壤均死亡。综合生理指标、寒害指数、保存率,10种受试植物的抗寒能力依次为：秋茄>桐花树>海漆>无瓣海桑>其它。生长能力依次为：无瓣海桑>拉关木>秋茄>桐花树>白骨壤。秋茄、桐花树、海漆和无瓣海桑可作为发展浙江人工红树林的优选树种。研究结果印证了全球变化背景下对浙江人工红树林造林树种引选的预测结果。
(3)浙江人工红树林造林树种对盐度的响应。盐度对红树植物光合作用的影响主要通过影响色素的合成、改变气孔运动、影响磷酸核酮糖羧化酶最大羧化效率和饱和光强下电子传递效率,以及影响光能捕获效率和电子传递量子效率等途径来实现。秋茄的盐度适生范围较无瓣海桑广,其在≦340 mM NaCl的盐度生境均能较好生长,170 mM NaCl是最佳盐度生境；无瓣海桑生境盐度应低于170 mMNaC。
(4)浙江人工红树林造林树种对潮汐浸淹的响应。以秋茄、无瓣海桑和桐花树为材料,研究了潮汐浸淹对其生长的影响。这三种红树植物均在中滩位保存率最高,秋茄和无瓣海桑在高滩位保存率高于低滩位,桐花树相反。以SOD活性、PRO含量、RE为参数分析受试红树植物对滩位的适应度(adp),相对于中滩位(adp=0),秋茄和桐花树在高滩位适应度降低(adp=—0.8204,adp=—0.7050),无瓣海桑稍有升高(adp=0.4795)；秋茄和无瓣海桑在低滩位适应度明显降低(adp= 3.4776,adp=—3.3934),而桐花树无变化(adp=—0.0988)。秋茄和无瓣海桑适应于中、高滩位,而桐花树适应于中、低滩位。
(5)不同滩位人工秋茄种群结构和生物量研究。滩位和林龄影响了种群特征,树高、冠幅和种群大小结构与树龄成显著正相关,与潮汐浸淹时间成显著负相关。种群密度与树龄成显著负相关,与潮汐浸淹时间成正相关。秋茄为基部多分枝型红树植物,通过测定其近地面分枝树干直径(D)与生物量,建立叶片(WL)、树干(WS)、地下部分(WB)及植株总生物量(WT)与树干直径D的异速生长式：WL=0.187×D1.855,Ws=O.267×D1.906,WB=4.6×D1.136,WT=3.614×D1.446。3年、5年和10年龄秋茄种群生物量分别为7.13±1.06、11.32±1.27和24.35±3.40 t/hm2,地下生物量占43.7%。
A large number of afforestation practices showed that temperature, salinity and intertidal position are the major limiting factors for mangrove forestation success. Therefore, study on the ecological responses of mangroves to temperature, salinity and intertidal position would provide theoretical basis and technical support to mangrove forestation in species and habitat selection. It has important location values in academic research on the study that ecological response of artificial mangroves to key environmental factors in Zhejiang Province, north area of mangrove introduction of China. In this study, field and greenhouse experiments were carried out to investigate to changes in plant growth, adaptation, survival, chilling injury index, photosynthesis and physiological parameters of 10 mangrove species, such as Kandelia obovata etc., in responding to key environmental factors, temperature, salinity and intertidal position, in order to clarify the adaptability of these mangroves to environmental factors though the methods of qualitative and quantitative analyses, the single index and the comprehensive analyses. Moreover, population structure and biomass were also investigated to provide theoretical basis for assessment of mangrove population dynamics and biomass of different ages in responding to intertidal position. The main results are as follows:
(1) Under the background of global climate change, the present paper forecasts for the introduction of artificial mangrove species in Zhejiang province:if the average temperature increased by 1.8℃, mangrove plants will achieve 3 species, K. obovata, Aegiceras corniculatum, distributed to Zhejiang Province. If the average temperature increased by 4℃, twelve mangrove species including to K. obovata, Avicennia marina, A. corniculatum, Sonneratia apetala etc, can be distributed to Zhejiang. It has important theoretical values on forecasts of Zhejiang artificial mangrove species selection.
(2) Response of Zhejiang artificial mangrove species to temperature: low temperature, especially extreme low temperature affected growth of mangroves. Relative electrolyte conductance (RE), malondiadehyde (MDA), proline (PRO) and superoxide dismutase (SOD) of experimental plants leaves increased with a decreasing temperature. After winter, plants of L. racemosa and A. marina were all dead (chilling injury index, Cii>3), some individuals of S. apetala were dead (Cii=2.76), while A. corniculatum, K. obovata and native species Ficuas concinna var. subsessilis were survival, with the similar values of Cii (1.08-1.14). After extreme low temperature (-4℃), the survival rate were K. obovata (77%)> A. corniculatum (73%)> E. agallocha (70%)> S. apetala (25%), whereas the other six mangrove species, including L. racemosa, B. gymnorhiza, B. sexangula var. rhynchopetala, R. stylosa, A. ilicifolius and A. marina, were all dead. As a result, cold tolerance of the experimental mangrove species as follows: K. obovata> A. corniculatum> E. agallocha> S. apetala > the other six mangrove species. Therefore, K. obovata, A. corniculatum, E. agallocha and S. apetala can be used as the main mangrove species in the artificial mangrove forestation in Zhejiang province, confirmed the context of mangrove species selection of Zhejiang artificial mangrove introduction.
(3) The effects of salinity on growth and photosynthesis were investigated in the two mangroves, S. apetala and K. obovata, under cultured conditions. The results showed that salinity affected plant photosynthesis by pigment synthesize, stomata changes, the values of the maximum rate of carboxylation by Rubisco and the PAR saturated electron transport rate, and the effective quantum yield etc. Kandelia obovata had a wider range of suitable salinity than S. apetala, which could grow well in≦340 Mm NaCl salinity habitats, with the optimum salinity of 170 Mm NaCl; whereas S. apetala had a strong low salinity preferences, with the salinity of its habitat should be less than 170 Mm NaCl.
(4) The effects of intertidal position on growth of K. obovata, S. apetala and A. corniculatum were tested, and the results were as followed: the three mangroves had the highest survival rate in middle intertidal position. The survival rate was higher in high than low intertidal position for K. obovata and S. apetala, and A. corniculatum opposite. The value of adaptation (adp) was estimated by the parameters of SOD, PRO and RE. The value of adp of K. obovata and A. corniculatum decreased,-0.8204 and -0.7050 for K. obovata and A. corniculatum, respectively, but increased for S. apetala (adp=0.4795) in high intertidal position compared to the adp value in middle intertidal position (adp =0). The adp values of K. obovata (adp=3.4776) and S. apetala (adp=-3.3934) in low intertidal position significantly decreased in comparison to middle intertidal position, while adp value did not change for A. corniculatum (adp=-0.0988). Therefore, K. obovata and S. apetala adapted to be planted in relative higher intertidal position, while A. corniculatum adapted to be planted in relative lower intertidal position.
(5) An experiment was conducted to investigate the population characteristics and biomass in artificial K. obovata mangrove in different intertidal position. The results showed that population characteristics such as height, canopy, and size structure positive relative to population age, but negative to intertidal position (tide-flooding time). Population density was negative to population age, while positive to tide-flooding time. Diameters of stems were used as variable for estimate plant biomass (leaf, WL; stem, Ws; belowground parts, WB; and total dry weight, WT). Allometric equations as followed: WL=0.187D1.855; Ws=0.267D1.906; WB=4.6D1.136; WT=3.614D1.446. Population biomass of 3-,5- and 10-year K. obovata was estimated at 7.13,11.32 and 24.35 t hm-2, with an average of 43.7% corresponding to belowground biomass.
(1)在全球气候变化背景下,对浙江红树林分布状况进行了预测。若年均温升温1.8℃,我国红树植物分布北界由福建福鼎向北推延至浙江舟山,分布到浙江的红树植物将达到3种,为秋茄、白骨壤(Avicennia marina)、桐花树。若升温4.0℃℃,我国红树植物分布北界将向北推延至南京,分布到浙江的红树植物将达到12种,为秋茄、白骨壤、桐花树、海漆(Excoecaria agallocha)、无瓣海桑等。该工作对浙江人工红树林造林树种的引选具有理论指导意义。
(2)浙江人工红树林造林树种对温度的响应。低温尤其是极端低温严重影响红树植物的生长。随着温度的降低,受试植物叶片相对电导率(RE)、脯氨酸含量(PRO)、超氧化物歧化酶活性(SOD)、丙二醛含量(MDA)逐渐增大。越冬后,拉关木(Laguncularia racemosa)和白骨壤全部死亡(寒害指数,Cii>3),无瓣海桑部分死亡(Cii=2.76),桐花树和秋茄的寒害指数与乡土植物无柄小叶榕(Ficuas concinna var. subsessilis)相似(1.08-1.14),无冻害现象。-4℃C极端低温过后,保存率依次为秋茄(77%)>桐花树(73%)>海漆(70%)>无瓣海桑(25%),拉关木、木榄(Bruguiera gymnorhiza)、尖瓣海莲(Bruguiera sexangula var. rhynchopetala)、红海榄(Rhizophora stylosa)、老鼠簕(Acanthus ilicifolius)、白骨壤均死亡。综合生理指标、寒害指数、保存率,10种受试植物的抗寒能力依次为：秋茄>桐花树>海漆>无瓣海桑>其它。生长能力依次为：无瓣海桑>拉关木>秋茄>桐花树>白骨壤。秋茄、桐花树、海漆和无瓣海桑可作为发展浙江人工红树林的优选树种。研究结果印证了全球变化背景下对浙江人工红树林造林树种引选的预测结果。
(3)浙江人工红树林造林树种对盐度的响应。盐度对红树植物光合作用的影响主要通过影响色素的合成、改变气孔运动、影响磷酸核酮糖羧化酶最大羧化效率和饱和光强下电子传递效率,以及影响光能捕获效率和电子传递量子效率等途径来实现。秋茄的盐度适生范围较无瓣海桑广,其在≦340 mM NaCl的盐度生境均能较好生长,170 mM NaCl是最佳盐度生境；无瓣海桑生境盐度应低于170 mMNaC。
(4)浙江人工红树林造林树种对潮汐浸淹的响应。以秋茄、无瓣海桑和桐花树为材料,研究了潮汐浸淹对其生长的影响。这三种红树植物均在中滩位保存率最高,秋茄和无瓣海桑在高滩位保存率高于低滩位,桐花树相反。以SOD活性、PRO含量、RE为参数分析受试红树植物对滩位的适应度(adp),相对于中滩位(adp=0),秋茄和桐花树在高滩位适应度降低(adp=—0.8204,adp=—0.7050),无瓣海桑稍有升高(adp=0.4795)；秋茄和无瓣海桑在低滩位适应度明显降低(adp= 3.4776,adp=—3.3934),而桐花树无变化(adp=—0.0988)。秋茄和无瓣海桑适应于中、高滩位,而桐花树适应于中、低滩位。
(5)不同滩位人工秋茄种群结构和生物量研究。滩位和林龄影响了种群特征,树高、冠幅和种群大小结构与树龄成显著正相关,与潮汐浸淹时间成显著负相关。种群密度与树龄成显著负相关,与潮汐浸淹时间成正相关。秋茄为基部多分枝型红树植物,通过测定其近地面分枝树干直径(D)与生物量,建立叶片(WL)、树干(WS)、地下部分(WB)及植株总生物量(WT)与树干直径D的异速生长式：WL=0.187×D1.855,Ws=O.267×D1.906,WB=4.6×D1.136,WT=3.614×D1.446。3年、5年和10年龄秋茄种群生物量分别为7.13±1.06、11.32±1.27和24.35±3.40 t/hm2,地下生物量占43.7%。
A large number of afforestation practices showed that temperature, salinity and intertidal position are the major limiting factors for mangrove forestation success. Therefore, study on the ecological responses of mangroves to temperature, salinity and intertidal position would provide theoretical basis and technical support to mangrove forestation in species and habitat selection. It has important location values in academic research on the study that ecological response of artificial mangroves to key environmental factors in Zhejiang Province, north area of mangrove introduction of China. In this study, field and greenhouse experiments were carried out to investigate to changes in plant growth, adaptation, survival, chilling injury index, photosynthesis and physiological parameters of 10 mangrove species, such as Kandelia obovata etc., in responding to key environmental factors, temperature, salinity and intertidal position, in order to clarify the adaptability of these mangroves to environmental factors though the methods of qualitative and quantitative analyses, the single index and the comprehensive analyses. Moreover, population structure and biomass were also investigated to provide theoretical basis for assessment of mangrove population dynamics and biomass of different ages in responding to intertidal position. The main results are as follows:
(1) Under the background of global climate change, the present paper forecasts for the introduction of artificial mangrove species in Zhejiang province:if the average temperature increased by 1.8℃, mangrove plants will achieve 3 species, K. obovata, Aegiceras corniculatum, distributed to Zhejiang Province. If the average temperature increased by 4℃, twelve mangrove species including to K. obovata, Avicennia marina, A. corniculatum, Sonneratia apetala etc, can be distributed to Zhejiang. It has important theoretical values on forecasts of Zhejiang artificial mangrove species selection.
(2) Response of Zhejiang artificial mangrove species to temperature: low temperature, especially extreme low temperature affected growth of mangroves. Relative electrolyte conductance (RE), malondiadehyde (MDA), proline (PRO) and superoxide dismutase (SOD) of experimental plants leaves increased with a decreasing temperature. After winter, plants of L. racemosa and A. marina were all dead (chilling injury index, Cii>3), some individuals of S. apetala were dead (Cii=2.76), while A. corniculatum, K. obovata and native species Ficuas concinna var. subsessilis were survival, with the similar values of Cii (1.08-1.14). After extreme low temperature (-4℃), the survival rate were K. obovata (77%)> A. corniculatum (73%)> E. agallocha (70%)> S. apetala (25%), whereas the other six mangrove species, including L. racemosa, B. gymnorhiza, B. sexangula var. rhynchopetala, R. stylosa, A. ilicifolius and A. marina, were all dead. As a result, cold tolerance of the experimental mangrove species as follows: K. obovata> A. corniculatum> E. agallocha> S. apetala > the other six mangrove species. Therefore, K. obovata, A. corniculatum, E. agallocha and S. apetala can be used as the main mangrove species in the artificial mangrove forestation in Zhejiang province, confirmed the context of mangrove species selection of Zhejiang artificial mangrove introduction.
(3) The effects of salinity on growth and photosynthesis were investigated in the two mangroves, S. apetala and K. obovata, under cultured conditions. The results showed that salinity affected plant photosynthesis by pigment synthesize, stomata changes, the values of the maximum rate of carboxylation by Rubisco and the PAR saturated electron transport rate, and the effective quantum yield etc. Kandelia obovata had a wider range of suitable salinity than S. apetala, which could grow well in≦340 Mm NaCl salinity habitats, with the optimum salinity of 170 Mm NaCl; whereas S. apetala had a strong low salinity preferences, with the salinity of its habitat should be less than 170 Mm NaCl.
(4) The effects of intertidal position on growth of K. obovata, S. apetala and A. corniculatum were tested, and the results were as followed: the three mangroves had the highest survival rate in middle intertidal position. The survival rate was higher in high than low intertidal position for K. obovata and S. apetala, and A. corniculatum opposite. The value of adaptation (adp) was estimated by the parameters of SOD, PRO and RE. The value of adp of K. obovata and A. corniculatum decreased,-0.8204 and -0.7050 for K. obovata and A. corniculatum, respectively, but increased for S. apetala (adp=0.4795) in high intertidal position compared to the adp value in middle intertidal position (adp =0). The adp values of K. obovata (adp=3.4776) and S. apetala (adp=-3.3934) in low intertidal position significantly decreased in comparison to middle intertidal position, while adp value did not change for A. corniculatum (adp=-0.0988). Therefore, K. obovata and S. apetala adapted to be planted in relative higher intertidal position, while A. corniculatum adapted to be planted in relative lower intertidal position.
(5) An experiment was conducted to investigate the population characteristics and biomass in artificial K. obovata mangrove in different intertidal position. The results showed that population characteristics such as height, canopy, and size structure positive relative to population age, but negative to intertidal position (tide-flooding time). Population density was negative to population age, while positive to tide-flooding time. Diameters of stems were used as variable for estimate plant biomass (leaf, WL; stem, Ws; belowground parts, WB; and total dry weight, WT). Allometric equations as followed: WL=0.187D1.855; Ws=0.267D1.906; WB=4.6D1.136; WT=3.614D1.446. Population biomass of 3-,5- and 10-year K. obovata was estimated at 7.13,11.32 and 24.35 t hm-2, with an average of 43.7% corresponding to belowground biomass.
引文
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