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叶片性状包括叶片的结构性状和功能性状,是由遗传因素和环境条件共同决定的。不同生境的植物叶片性状间的差异体现了植物对环境的适应性[1-2]。叶片重要的性状指标能反映出植物偿还叶片养分和干物质投资时间的快慢[3],如比叶面积较小的植物叶片会将能量和物质用来构建叶片而非投资到光合器官,因而含有更多的厚壁组织和细胞壁成分[4-6],这势必引起叶片碳同化能力的下调。然而,叶片碳的储存在一定程度上受控于氮、磷可获得的量[7-8],碳/氮(C/N)和碳/磷(C/P)的比值更是在一定程度上反映了植物的养分利用效率,氮/磷(N/P)比则可以作为对植物生长起限制性作用的养分元素的指示剂[9-10]。因此,研究植物叶片性状和养分化学计量与环境和人工经营干扰的关系,有助于理解植物对环境的适应机制及对资源的利用和分配过程等[11],具有重要的科学价值和生产指导意义。高节竹Phyllostachys prominens俗称钢鞭哺鸡竹,是优良的笋材兼用竹种,具有竹笋产量高、品质佳、加工性能好,竹材径级较大、材质坚硬,生态适应性强,地下鞭系粗壮、延伸生长能力强等特点,在浙江省杭州市、湖州市等地广为栽培。高节竹出笋期4月中下旬至5月中旬,较雷竹Phyllostachys violascens和毛竹Phyllostachys edulis出笋迟,由于竹笋消费疲劳的原因,高节竹林经济效益一直以来都表现不佳。为顺应市场对高品质竹笋的大量需求,根据高节竹的生物学和生态学特性,浙江省杭州市桐庐县高节竹主产区开展了高节竹林地覆土控鞭高品质竹笋栽培措施的规模化应用。覆土控鞭栽培措施实施后高节竹笋期推迟约15 d,竹笋外观品质、营养品质和食味品质明显改善,可食率提高,竹笋香甜味增加,酸涩味及粗糙度降低[12],深受上海、杭州等地的消费者欢迎,竹笋供不应求,竹林经营效益显著提高,推广规模日趋增大。高节竹林地覆土控鞭栽培后,林分结构可以通过留笋养竹和伐竹等人工调控措施来维持丰产结构要求,但竹林地下部分的生长环境发生了变化,这是否会对高节竹立竹叶片形态和养分化学计量特征产生影响,进而影响高节竹生长呢?为此,开展了覆土控鞭栽培2 a和不覆土栽培的高节竹林1~3年生立竹叶片主要形态指标和碳、氮、磷质量分数及化学计量比的比较研究,旨在为高节竹林高效可持续经营提供参考。
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由表 1可知:随立竹年龄增加,除比叶面积外,高节竹主要叶性因子均总体呈增加趋势,且2年生、3年生立竹显著高于1年生立竹(P<0.05),而前两者间并显著差异;比叶面积则相反,为1年生立竹显著高于2年生、3年生立竹,且后两者间仅略有差异(P>0.05)。
表 1 覆土控鞭栽培高节竹叶片形态特征
Table 1. Leaf traits of 1-3-years-old Phyllostachys prominens from experimental stand
处理 立竹年龄/a 叶长/cm 叶宽/cm 叶形指数 叶面积/cm2 单叶质量/g 比叶面积/(cm2·g-1) 叶片厚度/mm 覆土栽培 1 10.60 ± 0.56bA 2.03 ± 0.14bA 5.26 ± 0.29bA 15.44 ±1.28bA 0.21 ± 0.01bA 73.53 ±3.68aA 0.14 ± 0.01bA 2 16.80 ± 0.63aA 2.41 ± 0.21aA 6.96 ± 0.38aA 28.92 ±1.89aA 0.51 ± 0.03aA 56.56 ± 3.58bA 0.17 ± 0.02aA 3 17.29 ± 1.03aA 2.38 ± 0.15aA 7.26 ± 0.45aA 29.13 ±2.21aA 0.49 ± 0.03aA 58.93 ±3.64bA 0.16 ± 0.01aA 不覆土栽培 1 10.84 ± 1.01bA 2.05 ± 0.12bA 5.28 ± 0.42bA 15.56 ±1.03bA 0.22 ± 0.02bA 70.27 ±3.56aA 0.14 ± 0.01bA 2 16.54 ± 1.25aA 2.43 ± 0.21aA 6.81 ± 0.39aA 27.47 ±2.11aA 0.52 ± 0.04aA 53.07 ±2.54bA 0.18 ± 0.02aA 3 17.04 ± 0.98aA 2.45 ± 0.22aA 6.95 ± 0.43aA 28.64 ± 1.98aA 0.50 ± 0.03aA 57.28 ±3.21bA 0.18 ± 0.01aA 说明:大写字母不同表示相同年龄立竹不同处理间差异显著(P < 0.05),小写字母不同表示同一立竹年龄不同处理间差异显著(P < 0.05),相同字母表示差异不显著(P>0.05)。 覆土栽培2 a后,高节竹1年生立竹叶长稍有降低,而2~3年生立竹叶长有小幅度增加,1~3年生立竹叶宽均有小幅度下降,1~3年生立竹叶片叶长和叶宽的变化使叶形指数总体上有小幅度升高,叶长、叶宽和叶形指数覆土控鞭栽培和不覆土栽培竹林间均无显著差异(P>0.05);覆土栽培2 a后高节竹1~3年生立竹叶片单叶面积和比叶面积总体上均有一定幅度的升高,而单叶质量和叶片厚度总体上下降,单叶面积、单叶质量、比叶面积和叶片厚度覆土控鞭栽培和不覆土栽培竹林间也均无显著差异(P>0.05)。上述分析表明:虽然覆土控鞭栽培后高节竹的地下鞭系生长环境发生了明显的变化,特别是温度和土壤水分,但对不同年龄立竹的叶片形态特征并没有产生明显影响,体现出高节竹很强的生态适应性。
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随立竹年龄增加,高节竹叶片碳含质量分数总体升高,1年生立竹碳显著低于2年生和3年生立竹(P<0.05),且后两者间并无显著差异,而氮、磷总体下降,1年生立竹氮、磷显著高于2年生和3年生立竹,且后两者间亦无显著差异(P>0.05)(表 2)。覆土控鞭栽培2 a后,高节竹1~3年生立竹叶片碳质量分数均显著升高,增幅分别为6.62%,9.25%和6.09%,差异达显著水平(P<0.05);覆土栽培2 a后高节竹1~3年生立竹叶片氮质量分数也有不同幅度的升高,增幅分别为4.12%,2.51%和1.99%,而叶片磷质量分数均有一定幅度的下降,降幅分别为0.55%,2.89%和0.75%,但1~3年立竹叶片氮、磷质量分数覆土控鞭栽培与不覆土栽培高节竹林间均无显著差异(P>0.05)。可见,高节竹林覆土控鞭栽培后对叶片光合产物碳会产生明显的影响,竹林光合碳同化能力可能得到提高,但从土壤中吸收转运氮、磷养分并没有明显的变化。
表 2 覆土控鞭栽培高节竹叶片碳、氮、磷质量分数
Table 2. Leaf C, N, P content for 1-3-year-old Phyllostachys prominens of experimental stand
处理 立竹年龄/a 碳/(mg.g-1) 氮/(mg.g-1) 磷/(mg.g-1) 覆土栽培 1 456.51 ± 20.32 bA 23.51 ± 1.23aA 1.79 ± 0.12aA 2 502.09 ± 15.23aA 20.39 ± 1.54bA 1.34 ± 0.11bA 3 488.35 ± 18.23 aA 18.96 ± 1.09bA 1.32 ± 0.12bA 不覆土栽培 1 428.16 ± 9.68bB 22.58 ± 2.01aA 1.80 ± 0.14aA 2 476.05 ± 22.32 aB 19.89 ± 1.12bA 1.42 ± 0.11bA 3 460.32 ± 19.56aA 18.59 ± 1.07bA 1.33 ± 0.12bA 说明:大写字母不同表示相同年龄立竹不同处理间差异显著(P < 0.05),小写字母不同表示同一立竹年龄不同处理间差异显著(P < 0.05),相同字母表示差异不显著(P>0.05)。 -
由表 3可知:随立竹年龄增加,高节竹1~3年生立竹叶片碳/氮(C/N)比和碳/磷(C/P)比均总体呈升高变化趋势,2年生和3年生立竹叶片C/N和C/P匀显著高于1年生立竹(P<0.05),且2年生和3年生立竹的C/N和C/P并无显著差异(P>0.05);叶片氮/磷(N/P)比虽亦呈升高变化规律,但立竹年龄间差异不显著(P>0.05)。覆土控鞭栽培2 a后,高节竹1~3年生立竹叶片C/N和N/P均有所升高,增幅分别为2.43%,2.90%,3.99%和4.87%,5.55%,2.79%,但与不覆土栽培高节竹林比较并无显著差异,而高节竹林1~3年生立竹叶片C/P覆土栽培后均有显著提高,增幅分别为7.32%,8.32%和11.06%,覆土控鞭栽培和不覆土栽培高节竹林之间差异均达显著水平(P<0.05)。说明覆土控鞭栽培并没有影响到高节竹的土壤氮、磷养分吸收转运,而且一定程度上提高了高节竹的氮、磷养分利用效率。
表 3 覆土控鞭栽培高节竹叶片碳、氮、磷化学计量比
Table 3. The C, N, P stoichiometry for leaf of 1-3-year-old Phyllostachys prominens from experimental stand
处理 立竹年龄/a 碳/氮(C/N) 碳/磷(C/P) 氮/磷(N/P) 1 19.42 ± 1.43bA 255.10 ± 16.14bA 13.14 ± 0.84aA 覆土栽培 2 24.62 ± 2.11aA 373.65 ± 20.16aA 15.21 ± 1.16aA 3 25.75 ± 2.13aA 369.80 ± 22.12aA 14.36 ± 1.09aA 1 18.96 ± 1.11bA 237.68 ± 12.12bA 12.53 ± 1.02aA 不覆土栽培 2 23.93 ± 1.56aA 344.96 ± 15.32aA 14.41 ± 1.11aA 3 24.76 ± 2.01aA 332.60 ± 18.23aB 13.97 ± 0.98aA 说明:大写字母不同表示相同年龄立竹不同处理间差异显著(P < 0.05),小写字母不同表示同一立竹年龄不同处理间差异显著(P < 0.05),相同字母表示差异不显著(P>0.05)。
Leaf morphology and C, N, and P stoichiometry of Phyllostachys prominens under soil cover with rhizome controlling cultivation
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摘要: 高节竹Phyllostachys prominens是优良的笋材兼用竹种,生态适应性强。采取覆土控鞭经营措施能显著提高高节竹笋外观品质、营养品质和食味品质及竹林经济效益。为摸清覆土控鞭栽培对高节竹生长的影响,为高节竹林高效可持续经营提供参考,测定了覆土控鞭栽培与对照高节竹林1~3年生立竹叶片的主要叶性因子和碳(C),氮(N),磷(P)质量分数,分析高节竹叶片形态和养分化学计量特征对覆土控鞭栽培的响应规律。结果表明:覆土控鞭栽培2 a后,高节竹1~3年生立竹叶片变细长,单叶面积和比叶面积略有增大,单叶质量和叶片厚度稍有下降,叶片形态指标覆土控鞭栽培和不覆土栽培竹林间差异均不显著(P>0.05)。覆土控鞭栽培的高节竹林1~3年生立竹叶片碳质量分数显著升高(P < 0.05),氮略有升高(P>0.05),磷稍有下降,C/N和N/P均有所升高(P>0.05),而C/P显著升高(P < 0.05)。覆土控鞭栽培对高节竹叶片形态并未产生明显的影响,对高节竹光合碳同化能力和氮、磷养分利用效率有促进作用,可见高节竹林采取科学的覆土控鞭高品质竹笋栽培措施能达到高效可持续经营的目标。Abstract: To determine the effect of soil cover with rhizome controlling cultivation on growth of Phyllostachys prominens, an excellent shoot and timber used bamboo species ecologically well adapted to its habitat, and to provide guidance for sustainable bamboo management, leaf traits as well as the content of leaf carbon, nitrogen, and phosphorus for 1-3 year-old Ph. prominens of soil cover with rhizome controlling cultivation and normal cultivation were studied. Furthermore, leaf morphology and nutrient stoichiometry were also analyzed. Results showed that 1-3 year-old Ph. prominens leaf of soil cover with rhizome controlling cultivation stand tended to be narrow and long with leaf area and specific leaf area increasing (P>0.05); whereas, leaf mass and thickness decreased slightly(P>0.05). Between soil cover with rhizome controlling cultivation and normal cultivation, there were no differences (P>0.05) for the same above mentioned leaf traits. However, leaf C content greatly increased (P < 0.05), N content increased slightly (P>0.05), and P content decreased a little (P>0.05). Also, C/N and N/P increased slightly (P>0.05) with C/P being greatly enhanced (P < 0.05). These results indicated that soil cover with rhizome controlling cultivation resulted in slight variation of leaf morphological plasticity for Ph. prominens and showed that with photosynthetic C fixation and nutrient utilization efficiency increasing, high and stable production of superior quality bamboo shoots could be promoted.
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表 1 覆土控鞭栽培高节竹叶片形态特征
Table 1. Leaf traits of 1-3-years-old Phyllostachys prominens from experimental stand
处理 立竹年龄/a 叶长/cm 叶宽/cm 叶形指数 叶面积/cm2 单叶质量/g 比叶面积/(cm2·g-1) 叶片厚度/mm 覆土栽培 1 10.60 ± 0.56bA 2.03 ± 0.14bA 5.26 ± 0.29bA 15.44 ±1.28bA 0.21 ± 0.01bA 73.53 ±3.68aA 0.14 ± 0.01bA 2 16.80 ± 0.63aA 2.41 ± 0.21aA 6.96 ± 0.38aA 28.92 ±1.89aA 0.51 ± 0.03aA 56.56 ± 3.58bA 0.17 ± 0.02aA 3 17.29 ± 1.03aA 2.38 ± 0.15aA 7.26 ± 0.45aA 29.13 ±2.21aA 0.49 ± 0.03aA 58.93 ±3.64bA 0.16 ± 0.01aA 不覆土栽培 1 10.84 ± 1.01bA 2.05 ± 0.12bA 5.28 ± 0.42bA 15.56 ±1.03bA 0.22 ± 0.02bA 70.27 ±3.56aA 0.14 ± 0.01bA 2 16.54 ± 1.25aA 2.43 ± 0.21aA 6.81 ± 0.39aA 27.47 ±2.11aA 0.52 ± 0.04aA 53.07 ±2.54bA 0.18 ± 0.02aA 3 17.04 ± 0.98aA 2.45 ± 0.22aA 6.95 ± 0.43aA 28.64 ± 1.98aA 0.50 ± 0.03aA 57.28 ±3.21bA 0.18 ± 0.01aA 说明:大写字母不同表示相同年龄立竹不同处理间差异显著(P < 0.05),小写字母不同表示同一立竹年龄不同处理间差异显著(P < 0.05),相同字母表示差异不显著(P>0.05)。 表 2 覆土控鞭栽培高节竹叶片碳、氮、磷质量分数
Table 2. Leaf C, N, P content for 1-3-year-old Phyllostachys prominens of experimental stand
处理 立竹年龄/a 碳/(mg.g-1) 氮/(mg.g-1) 磷/(mg.g-1) 覆土栽培 1 456.51 ± 20.32 bA 23.51 ± 1.23aA 1.79 ± 0.12aA 2 502.09 ± 15.23aA 20.39 ± 1.54bA 1.34 ± 0.11bA 3 488.35 ± 18.23 aA 18.96 ± 1.09bA 1.32 ± 0.12bA 不覆土栽培 1 428.16 ± 9.68bB 22.58 ± 2.01aA 1.80 ± 0.14aA 2 476.05 ± 22.32 aB 19.89 ± 1.12bA 1.42 ± 0.11bA 3 460.32 ± 19.56aA 18.59 ± 1.07bA 1.33 ± 0.12bA 说明:大写字母不同表示相同年龄立竹不同处理间差异显著(P < 0.05),小写字母不同表示同一立竹年龄不同处理间差异显著(P < 0.05),相同字母表示差异不显著(P>0.05)。 表 3 覆土控鞭栽培高节竹叶片碳、氮、磷化学计量比
Table 3. The C, N, P stoichiometry for leaf of 1-3-year-old Phyllostachys prominens from experimental stand
处理 立竹年龄/a 碳/氮(C/N) 碳/磷(C/P) 氮/磷(N/P) 1 19.42 ± 1.43bA 255.10 ± 16.14bA 13.14 ± 0.84aA 覆土栽培 2 24.62 ± 2.11aA 373.65 ± 20.16aA 15.21 ± 1.16aA 3 25.75 ± 2.13aA 369.80 ± 22.12aA 14.36 ± 1.09aA 1 18.96 ± 1.11bA 237.68 ± 12.12bA 12.53 ± 1.02aA 不覆土栽培 2 23.93 ± 1.56aA 344.96 ± 15.32aA 14.41 ± 1.11aA 3 24.76 ± 2.01aA 332.60 ± 18.23aB 13.97 ± 0.98aA 说明:大写字母不同表示相同年龄立竹不同处理间差异显著(P < 0.05),小写字母不同表示同一立竹年龄不同处理间差异显著(P < 0.05),相同字母表示差异不显著(P>0.05)。 -
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