小时间尺度下东北小兴安岭和完达山地区鹿类冬季营养生态学研究
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摘要
2009年-2011年1月,作者采集了黑龙江省小兴安岭沾河林区的驼鹿和狍的粪便样本和完达山地区的马鹿粪便样本,运用降序对应分析方法,研究了取样期间驼鹿、狍和马鹿种群的主要植物组分。并整合在小兴安岭地区2006年1-3月和2007年1-3月的食性数据、生存所需的最重要的蛋白质和能量,以及影响到蛋白质消化的植物单宁含量,进行了响应曲面模型构建,研究了小兴安岭地区驼鹿、狍种群和完达山地区马鹿种群的营养策略。
2006,2007,2009和2010年1月作者对小兴安岭地区驼鹿、狍以及2009年和2010年1-2月完达山地区马鹿的雪尿样本进行了采集,并于2012年7-8月于北美阿拉斯加驼鹿研究中心和怀俄明州夏延山地动物园采集了11只驼鹿的纯尿液样本。在保护生理学范畴内,选用了尿液嘌呤衍生物以及尿液嘌呤衍生物与肌酐的比值作为适用于小时间尺度,且能够精确评估鹿类营养状态的指标。计算了马鹿在取样时间段内可消化干物质的摄入量和可代谢能摄入量,并通过构建人工神经网络模型这一计算生态学手段,筛选并优化了小时间尺度内对驼鹿和狍营养状态影响最大的气候因子,并得出了相应的数量关系,最后针对雪深,将时间尺度扩大到年间,讨论雪深对鹿类营养状态的影响。研究结果主要有以下4点：
(1)在小兴安岭地区,驼鹿和狍分别在2009年和2011年食物多样性达到最高,完达山地区马鹿在2009年也显现出了较高的食物多样性,根据最优取食理论,更高的食物多样性有可能意味着鹿类动物需要更多种植物维持其营养需求,但这种基于最优取食理论的推测是较为片面的,有时可能会导致较大偏差。
小兴安岭驼鹿种群2009年和2010年1月的主要食物组分为杨、柳、白桦和云杉。云杉的出现表明驼鹿在取样时间段内正在经受较为严重的营养限制。有所不同的是与驼鹿同域分布的狍种群的食物组分并没有很多针叶树种出现,在2009和2010年其主要食物组分包括杨、柳、白桦、榛和部分草本植物,在2011年其食物组分出现了一定的变化,蒙古栎作为另一种主要食物出现在狍种群的食物组分中,而相应的,杨、柳、白桦等植物出现相应的降低。狍种群的营养状况在取样时间段内可能稍好于驼鹿。完达山地区马鹿种群的食物组成与之前的研究出现较大的不同,马鹿在2009-2011年1-2月不再以榆、紫椴、榛和暴马丁香等植物为主要食物,转而以柳、白桦和杨树等植物为主要食物,一方面可能是作者所选的取样时间有所不同,马鹿在不同的时间段内极有可能采食不同的植物,另一方面则有可能是马鹿受环境影响,对营养物质的需求发生了变化。
(2)在对驼鹿和狍进行采食策略响应曲面建模时,为避免云杉相对较低的蛋白质含量和较高的单宁含量对驼鹿的营养策略响应曲面造成过大影响,即作者已知驼鹿会趋向于采食高单宁含量的植物,作者在建模时将云杉和冷杉排除在外,更倾向于了解在没有云杉影响时驼鹿是否依然趋向于采食高单宁含量的食物。并且作者对驼鹿和狍选取了相同的数学关系,以便发现二者可能存在的对冬季严酷多变环境不同的响应模式,建模结果均显著。
对于小兴安岭地区的驼鹿和完达山区的马鹿,作者认为二者会倾向于采食能量蛋白质和单宁处在一定范围内的食物,使自身的能量需求达到稳态,而这种稳态或临界值会随着环境的变化产生波动。小兴安岭地区的狍总体上则选择了高能量,高蛋白质,低单宁的营养策略,但在蛋白质和单宁含量达到特定区间时,也有出现对单宁的偏好的可能性。
(3)散养驼鹿尿液嘌呤衍生物测定结果表明,尿囊素在驼鹿尿液中的含量极低,野生驼鹿的尿液嘌呤衍生物含量甚至低于液相色谱的检测极限,这可能是驼鹿适应严寒氧化损伤而形成的降低尿酸氧化酶活性的对策。小兴安岭狍种群中也存在着尿囊素缺失的情况,作者推测这是与驼鹿相似的适应严寒的策略,但这个猜测还有待精确实验予以证实。而作者认为完达山马鹿由于处在较低纬度,取样时期内气候相对温和,并不需要这种抗氧化的保护机制,并未出现尿囊素缺失的情况。
对于小兴安岭的驼鹿和狍种群,尿液嘌呤衍生物的研究结果表明,在取样的2006,2007,2009和2010四年1月的取样日期内,驼鹿种群和狍种群都存在的营养不充足情况,其中2009年最为严重。这与通过食性和营养策略所得出的结论有所不同。
对于完达山地区的马鹿种群,根据已有针对雌性成年马鹿的数量关系,作者将马鹿雪尿尿囊素肌酐比值进行双侧截尾,将数值分布保留在雌性马鹿分布范围内,计算了在取样时间段,可消化干物质和可代谢能的摄入量,定量研究了马鹿冬季的营养物质摄入量,结果表明,在2009年和2010年,可消化干物质量摄入量均表现出供给不足,可代谢能摄入量在2010年满足了最低需求但在2009年有一部分个体摄入量不足。可见2009年1-2月对于马鹿也是较为严酷的一段时期。作者认为一定程度的补饲是必要的。
(4)在气候因子的相关性研究中并未发现较为统一的规律,因此作者通过构建人工神经网络模型,这一适合批量数据处理的基于非线性手段的方法,对小兴安岭的驼鹿和狍尿液嘌呤衍生物对气候因子变量的响应进行了筛选优化,结果显示,日最低温度和日平均风速是众多气候因子中对尿液嘌呤衍生物最为显著的因子,并求得了气候因子与尿液嘌呤衍生物的数量关系。
最后,相对较大的时间尺度下,雪深表现出对鹿类嘌呤衍生物和肌酐比值的均值有较为显著的影响,并建议雪深应该用于较长时间尺度的营养生态学研究中。
Moose and roe deer fecal samples were collected during January from2009-2011in the Lesser Khingan Mountains, as well as red deer fecal samples in Wanda Mountains in Heilongjiang Province. We analyzed major food component of moose, roe deer and red deer by using detrench correspondence analysis. Then, we integrated the food component data in January to March2006and2007in the same research area together with data on2009-2011and established response surface model.
In the mean time, we collected snow urine samples of moose and roe deer in January of2006,2007,2009and2010in the Lesser Khingan Mountains as well as red deer snow urine samples from Wanda Mountains in January and February of2009-2011. What's more, we collected11pure urine samples from Alaska Moose Research Center and Chenney Mountain Zoo. In the field of conservation physiology, we chose urinary purine derivatives and their ratio to creatinine for the fine time-scale analysis, which could estimate deer nutritional condition preciously. We accurately calculated the daily digestible dry matter intake and metabolizable energy intake of red deer distributing in the Wanda Mountains and used a computational ecology method-BP artificial neural network to select and optimize the most influential climate factor to purine derivatives to creatinine ratios. Then response surface model was established using the selected climate data and purine derivatives to creatinine ratios as independent factors and response factors respectively. Then we enlarged the research time scale to years and discussed the influence of snow on purine derivatives. Generally, there are four major aspects of results:
(1) In the Lesser Khingan Mountains, food diversity was the higest for moose in2009and roe deer in2011and in Wanda Mountains. According to the optimal forage theory, higher food diversity probability means that deers need more species to maintain their nutritional requirements, however, this statement could be biased based on the optimal foraging theory. Main food component of moose population distributing in the Lesser Khingan Mountains were poplar, willows, birch and spruce. Appearance of spruce showed a constrained nutritional condition in our research period. In contrast, the sympatric roe deer did not show a large component of needles in faces. In2009and2010main food component of roe deer were poplar, willows, birch, hazel and grass, while Mongolian oak became one of the main food components of roe deer in2011, and percentage of poplar, willows, birch became lower than2009and2010. This indicated that nutritional condition of roe deer might be better than moose in the Lesser Khingan Mountains. We found great difference between main food component in January and February of2009-2011and previous studies of red deer in Wanda Mountains. They do not select elm, amur linden and hazel as their main food but willows, birch and poplars. On one hand, this could be caused by different sampling time, on the other hand, the nutritional requirements changed under the influence of changing environmental condition.
(2) During the establishment of response surface model, as we have already know that spruce has high content of tannins and that moose in the Lesser Khingan Mountains select spruce as one of their main food, so in order to avoid the influence of spruce, we tending to know that whether moose still prefer food with high tannin content without the influence of spruce. Moreover, we used the same model for the sympatric moose and roe deer to discover different response pattern of the two species. Modeling building results were significant.
Moose distributing in the Lesser Khingan Mountains and red deer in the Wanda Mountains were tending to feed on food that have certain range of protein and tannins to reach a homostasis of their nutritional requirement, but this homostasis will fluctuate when environment changes. Generally, roe deer showed a preference of high protein and energy content food and avoidance of tannins, while still it is not difficult to figure out that when plant protein and tannins content achieve a certain range, roe deer showed a trend of preference to tannins.
Results of purine derivatives detection in pure urine from free ranging mooses in North America indicated that, allantoin content was extremely low, and allantoin content in snow urine from wild population distributing in the Lesser Khingan Mountains was even lower than the detection limit of high performance chromatography. The low content of allantoin in the Lesser Khingan Mountains was supposed to be a strategy of antioxidant of adapting to extremely cold winter by this might be the same strategy to moose selected by roe deer in the Lesser Khingan Mountains to resist cold weather. But this still need more accurate experiment before we could confirm this hypothesis. And there is no allantoin absence phenomenon in red deer population in the Wanda Mountains, which might because the red deer population distributing in a low latitude and do not need this conservation strategy.
During the sampling period in2006,2007,2009and2010, purine derivatives to creatinine ratios showed that both roe deer and moose populations were suffering from nutritional constrain, and the situation in2009is worse. This result was different from that we get from food component and food strategy analysis.
According to the exist mathematical relationships between allantoin to creatinine ratio and digestible dry matter intake and metabolizable energy intake in female adult red deers, we trimmed off part of the allantoin to creatinine ratio from both sides of data distribution to keep the left data could best reflect the condition of adult females in the Wanda Mountains. Then we calculated the digestible dry matter intake and metabolizable energy intake of red deer by solving equations. Results showed that digestible dry matter intake was in short supply in both2009and2010. Metabolizable energy intake meet the minimum requirement of red deer in2010but in2009part of the individuals was short in metabolizable energy intake supply. These demonstrated that January-February in2009is a relative serious time and some supplementary (4) Climate factors did not show a common regulation, though there was some correlations among factors, but we could not judge which have the greatest influence. Therefore, we used artificial neural network, which is one of the computational ecology methods used for nonlinear analysis and modeling, to select and optimize the climate factors that had the most influential effects. Results showed that daily minimum temperature and daily average temperature had the best prediction effects to purine derivatives to creatinine ratios. Then response surface model were built to describe the mathematical relationships of daily minimum temperature and daily average temperature and urinary purine derivatives to creatinine ratio. Surprisingly, snow depth did not have good prediction effect. Then we enlarged our research time scale to years we found that snow depth should be used in long time scale nutritional ecology research.
2006,2007,2009和2010年1月作者对小兴安岭地区驼鹿、狍以及2009年和2010年1-2月完达山地区马鹿的雪尿样本进行了采集,并于2012年7-8月于北美阿拉斯加驼鹿研究中心和怀俄明州夏延山地动物园采集了11只驼鹿的纯尿液样本。在保护生理学范畴内,选用了尿液嘌呤衍生物以及尿液嘌呤衍生物与肌酐的比值作为适用于小时间尺度,且能够精确评估鹿类营养状态的指标。计算了马鹿在取样时间段内可消化干物质的摄入量和可代谢能摄入量,并通过构建人工神经网络模型这一计算生态学手段,筛选并优化了小时间尺度内对驼鹿和狍营养状态影响最大的气候因子,并得出了相应的数量关系,最后针对雪深,将时间尺度扩大到年间,讨论雪深对鹿类营养状态的影响。研究结果主要有以下4点：
(1)在小兴安岭地区,驼鹿和狍分别在2009年和2011年食物多样性达到最高,完达山地区马鹿在2009年也显现出了较高的食物多样性,根据最优取食理论,更高的食物多样性有可能意味着鹿类动物需要更多种植物维持其营养需求,但这种基于最优取食理论的推测是较为片面的,有时可能会导致较大偏差。
小兴安岭驼鹿种群2009年和2010年1月的主要食物组分为杨、柳、白桦和云杉。云杉的出现表明驼鹿在取样时间段内正在经受较为严重的营养限制。有所不同的是与驼鹿同域分布的狍种群的食物组分并没有很多针叶树种出现,在2009和2010年其主要食物组分包括杨、柳、白桦、榛和部分草本植物,在2011年其食物组分出现了一定的变化,蒙古栎作为另一种主要食物出现在狍种群的食物组分中,而相应的,杨、柳、白桦等植物出现相应的降低。狍种群的营养状况在取样时间段内可能稍好于驼鹿。完达山地区马鹿种群的食物组成与之前的研究出现较大的不同,马鹿在2009-2011年1-2月不再以榆、紫椴、榛和暴马丁香等植物为主要食物,转而以柳、白桦和杨树等植物为主要食物,一方面可能是作者所选的取样时间有所不同,马鹿在不同的时间段内极有可能采食不同的植物,另一方面则有可能是马鹿受环境影响,对营养物质的需求发生了变化。
(2)在对驼鹿和狍进行采食策略响应曲面建模时,为避免云杉相对较低的蛋白质含量和较高的单宁含量对驼鹿的营养策略响应曲面造成过大影响,即作者已知驼鹿会趋向于采食高单宁含量的植物,作者在建模时将云杉和冷杉排除在外,更倾向于了解在没有云杉影响时驼鹿是否依然趋向于采食高单宁含量的食物。并且作者对驼鹿和狍选取了相同的数学关系,以便发现二者可能存在的对冬季严酷多变环境不同的响应模式,建模结果均显著。
对于小兴安岭地区的驼鹿和完达山区的马鹿,作者认为二者会倾向于采食能量蛋白质和单宁处在一定范围内的食物,使自身的能量需求达到稳态,而这种稳态或临界值会随着环境的变化产生波动。小兴安岭地区的狍总体上则选择了高能量,高蛋白质,低单宁的营养策略,但在蛋白质和单宁含量达到特定区间时,也有出现对单宁的偏好的可能性。
(3)散养驼鹿尿液嘌呤衍生物测定结果表明,尿囊素在驼鹿尿液中的含量极低,野生驼鹿的尿液嘌呤衍生物含量甚至低于液相色谱的检测极限,这可能是驼鹿适应严寒氧化损伤而形成的降低尿酸氧化酶活性的对策。小兴安岭狍种群中也存在着尿囊素缺失的情况,作者推测这是与驼鹿相似的适应严寒的策略,但这个猜测还有待精确实验予以证实。而作者认为完达山马鹿由于处在较低纬度,取样时期内气候相对温和,并不需要这种抗氧化的保护机制,并未出现尿囊素缺失的情况。
对于小兴安岭的驼鹿和狍种群,尿液嘌呤衍生物的研究结果表明,在取样的2006,2007,2009和2010四年1月的取样日期内,驼鹿种群和狍种群都存在的营养不充足情况,其中2009年最为严重。这与通过食性和营养策略所得出的结论有所不同。
对于完达山地区的马鹿种群,根据已有针对雌性成年马鹿的数量关系,作者将马鹿雪尿尿囊素肌酐比值进行双侧截尾,将数值分布保留在雌性马鹿分布范围内,计算了在取样时间段,可消化干物质和可代谢能的摄入量,定量研究了马鹿冬季的营养物质摄入量,结果表明,在2009年和2010年,可消化干物质量摄入量均表现出供给不足,可代谢能摄入量在2010年满足了最低需求但在2009年有一部分个体摄入量不足。可见2009年1-2月对于马鹿也是较为严酷的一段时期。作者认为一定程度的补饲是必要的。
(4)在气候因子的相关性研究中并未发现较为统一的规律,因此作者通过构建人工神经网络模型,这一适合批量数据处理的基于非线性手段的方法,对小兴安岭的驼鹿和狍尿液嘌呤衍生物对气候因子变量的响应进行了筛选优化,结果显示,日最低温度和日平均风速是众多气候因子中对尿液嘌呤衍生物最为显著的因子,并求得了气候因子与尿液嘌呤衍生物的数量关系。
最后,相对较大的时间尺度下,雪深表现出对鹿类嘌呤衍生物和肌酐比值的均值有较为显著的影响,并建议雪深应该用于较长时间尺度的营养生态学研究中。
Moose and roe deer fecal samples were collected during January from2009-2011in the Lesser Khingan Mountains, as well as red deer fecal samples in Wanda Mountains in Heilongjiang Province. We analyzed major food component of moose, roe deer and red deer by using detrench correspondence analysis. Then, we integrated the food component data in January to March2006and2007in the same research area together with data on2009-2011and established response surface model.
In the mean time, we collected snow urine samples of moose and roe deer in January of2006,2007,2009and2010in the Lesser Khingan Mountains as well as red deer snow urine samples from Wanda Mountains in January and February of2009-2011. What's more, we collected11pure urine samples from Alaska Moose Research Center and Chenney Mountain Zoo. In the field of conservation physiology, we chose urinary purine derivatives and their ratio to creatinine for the fine time-scale analysis, which could estimate deer nutritional condition preciously. We accurately calculated the daily digestible dry matter intake and metabolizable energy intake of red deer distributing in the Wanda Mountains and used a computational ecology method-BP artificial neural network to select and optimize the most influential climate factor to purine derivatives to creatinine ratios. Then response surface model was established using the selected climate data and purine derivatives to creatinine ratios as independent factors and response factors respectively. Then we enlarged the research time scale to years and discussed the influence of snow on purine derivatives. Generally, there are four major aspects of results:
(1) In the Lesser Khingan Mountains, food diversity was the higest for moose in2009and roe deer in2011and in Wanda Mountains. According to the optimal forage theory, higher food diversity probability means that deers need more species to maintain their nutritional requirements, however, this statement could be biased based on the optimal foraging theory. Main food component of moose population distributing in the Lesser Khingan Mountains were poplar, willows, birch and spruce. Appearance of spruce showed a constrained nutritional condition in our research period. In contrast, the sympatric roe deer did not show a large component of needles in faces. In2009and2010main food component of roe deer were poplar, willows, birch, hazel and grass, while Mongolian oak became one of the main food components of roe deer in2011, and percentage of poplar, willows, birch became lower than2009and2010. This indicated that nutritional condition of roe deer might be better than moose in the Lesser Khingan Mountains. We found great difference between main food component in January and February of2009-2011and previous studies of red deer in Wanda Mountains. They do not select elm, amur linden and hazel as their main food but willows, birch and poplars. On one hand, this could be caused by different sampling time, on the other hand, the nutritional requirements changed under the influence of changing environmental condition.
(2) During the establishment of response surface model, as we have already know that spruce has high content of tannins and that moose in the Lesser Khingan Mountains select spruce as one of their main food, so in order to avoid the influence of spruce, we tending to know that whether moose still prefer food with high tannin content without the influence of spruce. Moreover, we used the same model for the sympatric moose and roe deer to discover different response pattern of the two species. Modeling building results were significant.
Moose distributing in the Lesser Khingan Mountains and red deer in the Wanda Mountains were tending to feed on food that have certain range of protein and tannins to reach a homostasis of their nutritional requirement, but this homostasis will fluctuate when environment changes. Generally, roe deer showed a preference of high protein and energy content food and avoidance of tannins, while still it is not difficult to figure out that when plant protein and tannins content achieve a certain range, roe deer showed a trend of preference to tannins.
Results of purine derivatives detection in pure urine from free ranging mooses in North America indicated that, allantoin content was extremely low, and allantoin content in snow urine from wild population distributing in the Lesser Khingan Mountains was even lower than the detection limit of high performance chromatography. The low content of allantoin in the Lesser Khingan Mountains was supposed to be a strategy of antioxidant of adapting to extremely cold winter by this might be the same strategy to moose selected by roe deer in the Lesser Khingan Mountains to resist cold weather. But this still need more accurate experiment before we could confirm this hypothesis. And there is no allantoin absence phenomenon in red deer population in the Wanda Mountains, which might because the red deer population distributing in a low latitude and do not need this conservation strategy.
During the sampling period in2006,2007,2009and2010, purine derivatives to creatinine ratios showed that both roe deer and moose populations were suffering from nutritional constrain, and the situation in2009is worse. This result was different from that we get from food component and food strategy analysis.
According to the exist mathematical relationships between allantoin to creatinine ratio and digestible dry matter intake and metabolizable energy intake in female adult red deers, we trimmed off part of the allantoin to creatinine ratio from both sides of data distribution to keep the left data could best reflect the condition of adult females in the Wanda Mountains. Then we calculated the digestible dry matter intake and metabolizable energy intake of red deer by solving equations. Results showed that digestible dry matter intake was in short supply in both2009and2010. Metabolizable energy intake meet the minimum requirement of red deer in2010but in2009part of the individuals was short in metabolizable energy intake supply. These demonstrated that January-February in2009is a relative serious time and some supplementary (4) Climate factors did not show a common regulation, though there was some correlations among factors, but we could not judge which have the greatest influence. Therefore, we used artificial neural network, which is one of the computational ecology methods used for nonlinear analysis and modeling, to select and optimize the climate factors that had the most influential effects. Results showed that daily minimum temperature and daily average temperature had the best prediction effects to purine derivatives to creatinine ratios. Then response surface model were built to describe the mathematical relationships of daily minimum temperature and daily average temperature and urinary purine derivatives to creatinine ratio. Surprisingly, snow depth did not have good prediction effect. Then we enlarged our research time scale to years we found that snow depth should be used in long time scale nutritional ecology research.
引文
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