攀援植物爬山虎生长特性及其在高陡岩面植被恢复中的应用
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摘要
攀援植物是一类不能直立，需要通过主茎缠绕或攀附器官攀附支持物生长的植物总称，这类植物是热带和亚热带森林中重要物种，分布广泛，是药材、食品原料和景观绿化植物重要来源；同时攀援植物也是杂草控制主要种类，其缠绕攀附的生长行为也危害其他植物的生长。研究攀援植物的生长特点和养分吸收规律，发现攀援植物生长的限制因子，将有助于合理掌握攀援植物在生态环境中的应用尺度。本文以典型的亚热带多年生落叶木本攀援植物爬山虎(Parthenocissustricuspidata(Sieb．et Zucc))为研究对象，对自然生长爬山虎的营养元素分布和季节变化规律，氮磷钾养分、支持物、光照、高温和水分等环境条件对攀援植物的生长的影响以及其在高陡岩面的植被恢复的应用等方面进行了研究，结果如下：
1．自然生长爬山虎地上部养分分布不仅与器官有关，而且受器官所在部位及季节变化的影响；叶片的养分含量显著高于茎和叶柄；植株上部器官的养分含量低于中部和下部器官的养分含量；而在不同部位和不同器官的各类营养元素随季节变化有不同的变化趋势。爬山虎各器官的养分含量次序有明显不同，叶片营养元素的含量顺序是N＞Ca＞K＞Mg＞P＞Fe＞Mn＞B＞Zn＞Cu，叶柄的营养元素的含量顺序是N＞Ca＞K＞Mg＞P＞Fe＞B＞Mn＞Zn＞Cu，茎的营养元素的含量顺序是N＞Ca＞K＞P＞Mg＞Fe＞Mn＞B＞Zn＞Cu。爬山虎植株各营养元素的转移率以N和K最高，达到75％以上；Mn的转移率最低，仅为10．6％。
2．供氮水平的提高能增加爬山虎的生物量，爬山虎能够通过主茎和侧枝的生长形态的变化适应不同的氮素水平。植株茎、叶和根的氮含量和氮累积量也随着供氮水平的提高而增加，叶是整个植株的氮贮存的主要器官，叶的氮累积量达到整个植株的60％-70％；供氮水平的增加降低了爬山虎的氮利用率，提高了磷钾的利用率，并且磷的利用率高于氮钾的利用率。
3．供磷水平增加，爬山虎主茎的生长速度和生物量显著提高，但供磷水平过高后(大于0．06 g．L~(-1))对爬山虎的生长影响不显著；低磷水平显著影响爬山虎的根构型的变化，导致其爬山虎根系生物量占总生物量的比例增大；爬山虎在0．03 g．L~(-1)处理的总根长、根表面积、根系体积、根尖数和侧根数显著高于其他处理；随着供磷水平的增加，爬山虎根茎叶的磷含量和叶根氮含量有增加趋势，但降低了根茎叶的钾含量，而爬山虎的茎含氮量随供磷水平的增加先增加后减少；当供磷水平高于0．03 g．L~(-1)，对爬山虎根茎叶的氮磷钾的含量的影响不显著，表明爬山虎适应低磷养分供应；随着供磷水平的增加，对爬山虎的磷养分利用率影响不显著，但能提高爬山虎钾的养分利用率，减少氮的养分利用率。
4．供钾水平的增加，能增加爬山虎的生物量和主茎长度；过多的钾供应对生物量变化影响不显著，但调节各器官的生物量分配比例，主要表现为主茎生长受到抑制，增加叶在植株体内的生物量的分配比例；当供钾水平增加后，爬山虎植株体内的钾含量和钾累积量都显著增加；当供钾处理达到一定值后(0．18 g．L~(-1)处理)，爬山虎的钾含量和钾累积量没有显著提高，这表明爬山虎是对钾易满足型植物；供钾水平增加对爬山虎根和茎的氮含量影响不显著，能提高爬山虎叶的氮含量；对爬山虎茎的磷含量影响不显著，但降低叶根的磷含量；当供钾处理达到一定值后(0．18 g．L~(-1)处理)，爬山虎的根茎叶的氮磷钾含量没有显著变化。随着供钾水平的增加，各处理的爬山虎的氮利用率有降低趋势，磷养分利用率有升高趋势，但差异不显著；而钾的养分利用率随供钾水平的提高而显著减少。
5．匍匐生长的爬山虎植株主要通过增大主茎生物量和主茎长度而适应生长环境，而垂直攀援生长的爬山虎植株表现出较强的分枝能力，可以通过分枝茎的伸展以扩展生长空间。随着支持物角度的增加，爬山虎植株的主茎长度、主茎直径有减小的趋势，而分枝和分枝总长有增加的趋势。
6．适当遮荫有利于促进爬山虎的主茎纵向生长和分支的横向生长；但遮荫强度过大显著减少爬山虎生物量；遮荫强度不同影响爬山虎的各器官生物量分配，低遮荫条件有助于叶生物量积累，高遮荫导致爬山虎以茎生物量积累为主。
7．随着40℃高温时间持续，爬山虎叶片的相对含水量和蛋白质含量显著降低，而CMS和MDA显著升高；抗氧化酶(SOD、CAT和POD)先升高后降低。而高温持续第2天后，爬山虎的净光合值快速下降。
8．增加土壤水分能显著增加爬山虎的植株生物量，较高的土壤水分供应有助于爬山虎叶生物量分配，而较低的土壤水分有助于根生物量分配。土壤水分减少能使爬山虎叶片的气孔导度gs、蒸腾速率E和净光合值P_N显著下降。土壤水分变化，对爬山虎根茎叶的磷和钾含量影响较小，但显著影响爬山虎氮的含量。在不同的土壤水分条件下，爬山虎叶片保持了低的钾含量(低于10 g．kg~(-1))，有助于限制叶片气孔开度，这可能是爬山虎这种茎细叶茂结构适应水分环境的重要机理。土壤水分增加，对爬山虎的磷和钾的养分利用率影响不显著，但显著影响爬山虎的氮的养分利用率。
9．爬山虎在岩石坡面有较高的成活率和生长速度，可以作为先锋植物用于高陡岩面的植被恢复；将爬山虎结合岩面钻孔种植的方法可以作为高陡岩面植被恢复的技术之一，并优先在多雨地区和庇荫稳定的坡面使用。而使用压缩型生长基质(生态块)种植爬山虎有助于其在岩石坡面生长。
The climbing plants is an important species of tropical and subtropical forests,and canprovide deep green cover to many objects,rapidly climbing to supports by means of tendrils oradhesive disks for not standing.They are not only main raw material of medicine and food butalso planted widely for landscaping,soil erosion control and revegation.But sometimes,theclimbing plants act as weed to damage trees,shrubs and flower beds for windly growth anddifficultly control.So learning the growth features and uptake nutrients law of the climbing plantsand finding the limiting factor to growth will contribute to have reasonable application of them inenvironment.
Parthenocissus tricuspidata (Sieb.et Zucc) is a woody deciduous climbing plant of thefamily Vitaceae that has been widely planted for city green in china and play an important role inthe envinment management.But it is little known about its growth characteristics.This paperrepresented that the change of nutrients trend and distribution with the change of season,the N,P,K uptake characteristic,the effect of support angles,shaping,high temperature,soil moisture ongrowth,as well as revegetation of high and slope ofP.tricuspidata.The results were as follows:
1.With the season change,the concentration of the macronutirents N,P,K,Ca,Mg and themicronutrients Fe,Mn,B,Cu,Zn in different plant parts of natural grown P.tricuspidata haddifferent change trends.The nutrient concentration of leaf was higher than stem and leafstalk.Thetissue nutrient concentration of upside part of the plant was less than of middle part and downsidepart.Nutrient elements of different tissue had different concentration order:in leaf,N＞Ca＞K＞Mg＞P＞Fe＞Mn＞B＞Zn＞Cu;in leafstalk,N＞Ca＞K＞Mg＞P＞Fe＞B3＞Mn＞Zn＞Cu;in stem,N＞Ca＞K＞P＞Mg＞Fe＞Mn＞B＞Zn＞Cu.The N and K was the highest retranslocation rate among allnutrient elements in P.tricuspidata,exceeded 75%,and the Mn retranslocation rate was lowerthan other nutrient elements,only 10.6%.
2.Plant branches were smaller and shorter in size under lower nitrogen rates than higher rates.High nitrogen rates also significantly promoted early branching of P.tricuspidata seedlings andlowered branching positions on the main stems.Supplying high nitrogen to P.tricuspidatasignificantly increased plant biomass and nitrogen content in root,leaf and stem.Compared tostem and root,leaf had highest biomass,accounting for 50% of plant total biomass.Leaf was alsothe main tissue for nitrogen accumulation,accounting for 60% to 70% of total accumulatednitrogen.
3.Increasing phosphorus rates could increase the biomass and main stem length of P.tricuspidata,but high phosphorus rate (＞0.06 g.L~(-1)) did not influence significantly the growth of P.tricuspidata.There was significant variation of root style and biomass under lower phosphorusrate.The total root length,root surface area,root volume,root tips and forks under 0.03 g.L~(-1) Prate treatment was higher than other treatments.With phosphorus rates increasing,the tissuephosphorus concentration and leaf and root nitrogen concentration of P.tricuspidata had an increase trend while the tissue potassium concentration was reduced,but the stem nitrogenconcentration increased first,and then decreased later.Higher phosphorus rates (0.03 g.L~(-1))had noobvious effect on nitrogen,phosphorus and potassium concentration of stems,leaves and roots,and indicated that P.tricuspidata could adapt to lower phosphorus availability.Increasedphosphorus rates can increase potassium use efficiency and decrease nitrogen use efficiency,butno obviously influence on phosphorus use efficiency.
4.Increased potassium rates can increase the biomass and main stem length ofP.tricuspidata.However,higher potassium did not affect the biomass,but adjust the biomass partition.Leafgrowth constituted the more biomass proportion while the stem biomass was reduced under higherpotassium rates condition.The tissue potassium concentration and potassium content wereincreased with the increasing potassium rate.But when the potassium rate exceeded 0.18 g.L~(-1),theincrement of the tissue potassium concentration and potassium content was not significant.It issuggested that P.tricuspidata could adapt to lower potassium availability.Increased potassiumrates significantly increased the leaf nitrogen concentration,and reduced the leaf and rootphosphorus concentration,but not affected the root and stem nitrogen concentration and thestem phosphorus concentration.Higher potassium rates (＞0.18 g.L~(-1)) did not greatly influencethe tissue nutrient concentration.With the potassium rates increasing,potassium use efficiencydecreased significantly,and nitrogen use efficiency showed a falling trend while phosphorus useefficiency had a rising trend,but not significantly different.
5.The supports angles influenced the morphology and growth of P.tricuspidata.Under nosupporting condition,the biomass and main stem length of P.tricuspidata wasincreased.Undervertical angle condition,P.tricuspidata showed strong branch ability to extend growth space.When the support angles increased,the stem length and diameter of P.tricuspidata decreasedwhile the the branch number and total branch length inceased obviously.
6.Different shading measures obviously affected growth and biomass distribution of P.tricuspidata.Proper shading measures was good for developing the growth of main stem andbranches,and low shading increased the leaf biomass while high shading leaded to stem biomassincrease.
7.Under the high temperatures 40℃stress,the leaf relative water content (RWC),dissoluble protein content was significantly decreased but the cell membrane thermostability(CMS) and MDA were increased.Superoxide dismutase (SOD),catalase (CAT),peroxidase (POD)activities in leaf increased obviously at early stage of high temperature stress,then decreasedrapidly.After high temperatures stress for two days,the Net photosynthetic rate (P_N) was sharplyfalling.
8.Biomass in P.tricuspidata was increased with the enhancing of water supply.However,the seedlings grown under higher water supplies invested more biomass in leaf (in terms of leafnumber and leaf area),and root growth constituted the largest proportion of the biomass underlower water supplies.Decreased water supplies significantly reduced the Net photosynthetic rate (P_N),stomatal conductance (gs) and transpiration rate (E).Different water supplies greatlyinfluenced the tissue N concentration than P and K concentration.P.tricuspidata had a low Kconcentration (below 10 g.kg~(-1)) in all its plant tissues,which may allow the plants to limit itsstomatal opening and reduce its water loss.Water supplies had significantly affected N-useefficiency,but did not affect P and K-use efficiency.
9.P.tricuspidata had higher survival and growth rates,and showed well adaptability todisadvantageous growing conditions of rock slope so that it could be used as pioneer plant speciesto revegetation of rock slopes.The compress substrates can impove the biomass and growth rateof P.tricuspidata planting in rock slope.
1．自然生长爬山虎地上部养分分布不仅与器官有关，而且受器官所在部位及季节变化的影响；叶片的养分含量显著高于茎和叶柄；植株上部器官的养分含量低于中部和下部器官的养分含量；而在不同部位和不同器官的各类营养元素随季节变化有不同的变化趋势。爬山虎各器官的养分含量次序有明显不同，叶片营养元素的含量顺序是N＞Ca＞K＞Mg＞P＞Fe＞Mn＞B＞Zn＞Cu，叶柄的营养元素的含量顺序是N＞Ca＞K＞Mg＞P＞Fe＞B＞Mn＞Zn＞Cu，茎的营养元素的含量顺序是N＞Ca＞K＞P＞Mg＞Fe＞Mn＞B＞Zn＞Cu。爬山虎植株各营养元素的转移率以N和K最高，达到75％以上；Mn的转移率最低，仅为10．6％。
2．供氮水平的提高能增加爬山虎的生物量，爬山虎能够通过主茎和侧枝的生长形态的变化适应不同的氮素水平。植株茎、叶和根的氮含量和氮累积量也随着供氮水平的提高而增加，叶是整个植株的氮贮存的主要器官，叶的氮累积量达到整个植株的60％-70％；供氮水平的增加降低了爬山虎的氮利用率，提高了磷钾的利用率，并且磷的利用率高于氮钾的利用率。
3．供磷水平增加，爬山虎主茎的生长速度和生物量显著提高，但供磷水平过高后(大于0．06 g．L~(-1))对爬山虎的生长影响不显著；低磷水平显著影响爬山虎的根构型的变化，导致其爬山虎根系生物量占总生物量的比例增大；爬山虎在0．03 g．L~(-1)处理的总根长、根表面积、根系体积、根尖数和侧根数显著高于其他处理；随着供磷水平的增加，爬山虎根茎叶的磷含量和叶根氮含量有增加趋势，但降低了根茎叶的钾含量，而爬山虎的茎含氮量随供磷水平的增加先增加后减少；当供磷水平高于0．03 g．L~(-1)，对爬山虎根茎叶的氮磷钾的含量的影响不显著，表明爬山虎适应低磷养分供应；随着供磷水平的增加，对爬山虎的磷养分利用率影响不显著，但能提高爬山虎钾的养分利用率，减少氮的养分利用率。
4．供钾水平的增加，能增加爬山虎的生物量和主茎长度；过多的钾供应对生物量变化影响不显著，但调节各器官的生物量分配比例，主要表现为主茎生长受到抑制，增加叶在植株体内的生物量的分配比例；当供钾水平增加后，爬山虎植株体内的钾含量和钾累积量都显著增加；当供钾处理达到一定值后(0．18 g．L~(-1)处理)，爬山虎的钾含量和钾累积量没有显著提高，这表明爬山虎是对钾易满足型植物；供钾水平增加对爬山虎根和茎的氮含量影响不显著，能提高爬山虎叶的氮含量；对爬山虎茎的磷含量影响不显著，但降低叶根的磷含量；当供钾处理达到一定值后(0．18 g．L~(-1)处理)，爬山虎的根茎叶的氮磷钾含量没有显著变化。随着供钾水平的增加，各处理的爬山虎的氮利用率有降低趋势，磷养分利用率有升高趋势，但差异不显著；而钾的养分利用率随供钾水平的提高而显著减少。
5．匍匐生长的爬山虎植株主要通过增大主茎生物量和主茎长度而适应生长环境，而垂直攀援生长的爬山虎植株表现出较强的分枝能力，可以通过分枝茎的伸展以扩展生长空间。随着支持物角度的增加，爬山虎植株的主茎长度、主茎直径有减小的趋势，而分枝和分枝总长有增加的趋势。
6．适当遮荫有利于促进爬山虎的主茎纵向生长和分支的横向生长；但遮荫强度过大显著减少爬山虎生物量；遮荫强度不同影响爬山虎的各器官生物量分配，低遮荫条件有助于叶生物量积累，高遮荫导致爬山虎以茎生物量积累为主。
7．随着40℃高温时间持续，爬山虎叶片的相对含水量和蛋白质含量显著降低，而CMS和MDA显著升高；抗氧化酶(SOD、CAT和POD)先升高后降低。而高温持续第2天后，爬山虎的净光合值快速下降。
8．增加土壤水分能显著增加爬山虎的植株生物量，较高的土壤水分供应有助于爬山虎叶生物量分配，而较低的土壤水分有助于根生物量分配。土壤水分减少能使爬山虎叶片的气孔导度gs、蒸腾速率E和净光合值P_N显著下降。土壤水分变化，对爬山虎根茎叶的磷和钾含量影响较小，但显著影响爬山虎氮的含量。在不同的土壤水分条件下，爬山虎叶片保持了低的钾含量(低于10 g．kg~(-1))，有助于限制叶片气孔开度，这可能是爬山虎这种茎细叶茂结构适应水分环境的重要机理。土壤水分增加，对爬山虎的磷和钾的养分利用率影响不显著，但显著影响爬山虎的氮的养分利用率。
9．爬山虎在岩石坡面有较高的成活率和生长速度，可以作为先锋植物用于高陡岩面的植被恢复；将爬山虎结合岩面钻孔种植的方法可以作为高陡岩面植被恢复的技术之一，并优先在多雨地区和庇荫稳定的坡面使用。而使用压缩型生长基质(生态块)种植爬山虎有助于其在岩石坡面生长。
The climbing plants is an important species of tropical and subtropical forests,and canprovide deep green cover to many objects,rapidly climbing to supports by means of tendrils oradhesive disks for not standing.They are not only main raw material of medicine and food butalso planted widely for landscaping,soil erosion control and revegation.But sometimes,theclimbing plants act as weed to damage trees,shrubs and flower beds for windly growth anddifficultly control.So learning the growth features and uptake nutrients law of the climbing plantsand finding the limiting factor to growth will contribute to have reasonable application of them inenvironment.
Parthenocissus tricuspidata (Sieb.et Zucc) is a woody deciduous climbing plant of thefamily Vitaceae that has been widely planted for city green in china and play an important role inthe envinment management.But it is little known about its growth characteristics.This paperrepresented that the change of nutrients trend and distribution with the change of season,the N,P,K uptake characteristic,the effect of support angles,shaping,high temperature,soil moisture ongrowth,as well as revegetation of high and slope ofP.tricuspidata.The results were as follows:
1.With the season change,the concentration of the macronutirents N,P,K,Ca,Mg and themicronutrients Fe,Mn,B,Cu,Zn in different plant parts of natural grown P.tricuspidata haddifferent change trends.The nutrient concentration of leaf was higher than stem and leafstalk.Thetissue nutrient concentration of upside part of the plant was less than of middle part and downsidepart.Nutrient elements of different tissue had different concentration order:in leaf,N＞Ca＞K＞Mg＞P＞Fe＞Mn＞B＞Zn＞Cu;in leafstalk,N＞Ca＞K＞Mg＞P＞Fe＞B3＞Mn＞Zn＞Cu;in stem,N＞Ca＞K＞P＞Mg＞Fe＞Mn＞B＞Zn＞Cu.The N and K was the highest retranslocation rate among allnutrient elements in P.tricuspidata,exceeded 75%,and the Mn retranslocation rate was lowerthan other nutrient elements,only 10.6%.
2.Plant branches were smaller and shorter in size under lower nitrogen rates than higher rates.High nitrogen rates also significantly promoted early branching of P.tricuspidata seedlings andlowered branching positions on the main stems.Supplying high nitrogen to P.tricuspidatasignificantly increased plant biomass and nitrogen content in root,leaf and stem.Compared tostem and root,leaf had highest biomass,accounting for 50% of plant total biomass.Leaf was alsothe main tissue for nitrogen accumulation,accounting for 60% to 70% of total accumulatednitrogen.
3.Increasing phosphorus rates could increase the biomass and main stem length of P.tricuspidata,but high phosphorus rate (＞0.06 g.L~(-1)) did not influence significantly the growth of P.tricuspidata.There was significant variation of root style and biomass under lower phosphorusrate.The total root length,root surface area,root volume,root tips and forks under 0.03 g.L~(-1) Prate treatment was higher than other treatments.With phosphorus rates increasing,the tissuephosphorus concentration and leaf and root nitrogen concentration of P.tricuspidata had an increase trend while the tissue potassium concentration was reduced,but the stem nitrogenconcentration increased first,and then decreased later.Higher phosphorus rates (0.03 g.L~(-1))had noobvious effect on nitrogen,phosphorus and potassium concentration of stems,leaves and roots,and indicated that P.tricuspidata could adapt to lower phosphorus availability.Increasedphosphorus rates can increase potassium use efficiency and decrease nitrogen use efficiency,butno obviously influence on phosphorus use efficiency.
4.Increased potassium rates can increase the biomass and main stem length ofP.tricuspidata.However,higher potassium did not affect the biomass,but adjust the biomass partition.Leafgrowth constituted the more biomass proportion while the stem biomass was reduced under higherpotassium rates condition.The tissue potassium concentration and potassium content wereincreased with the increasing potassium rate.But when the potassium rate exceeded 0.18 g.L~(-1),theincrement of the tissue potassium concentration and potassium content was not significant.It issuggested that P.tricuspidata could adapt to lower potassium availability.Increased potassiumrates significantly increased the leaf nitrogen concentration,and reduced the leaf and rootphosphorus concentration,but not affected the root and stem nitrogen concentration and thestem phosphorus concentration.Higher potassium rates (＞0.18 g.L~(-1)) did not greatly influencethe tissue nutrient concentration.With the potassium rates increasing,potassium use efficiencydecreased significantly,and nitrogen use efficiency showed a falling trend while phosphorus useefficiency had a rising trend,but not significantly different.
5.The supports angles influenced the morphology and growth of P.tricuspidata.Under nosupporting condition,the biomass and main stem length of P.tricuspidata wasincreased.Undervertical angle condition,P.tricuspidata showed strong branch ability to extend growth space.When the support angles increased,the stem length and diameter of P.tricuspidata decreasedwhile the the branch number and total branch length inceased obviously.
6.Different shading measures obviously affected growth and biomass distribution of P.tricuspidata.Proper shading measures was good for developing the growth of main stem andbranches,and low shading increased the leaf biomass while high shading leaded to stem biomassincrease.
7.Under the high temperatures 40℃stress,the leaf relative water content (RWC),dissoluble protein content was significantly decreased but the cell membrane thermostability(CMS) and MDA were increased.Superoxide dismutase (SOD),catalase (CAT),peroxidase (POD)activities in leaf increased obviously at early stage of high temperature stress,then decreasedrapidly.After high temperatures stress for two days,the Net photosynthetic rate (P_N) was sharplyfalling.
8.Biomass in P.tricuspidata was increased with the enhancing of water supply.However,the seedlings grown under higher water supplies invested more biomass in leaf (in terms of leafnumber and leaf area),and root growth constituted the largest proportion of the biomass underlower water supplies.Decreased water supplies significantly reduced the Net photosynthetic rate (P_N),stomatal conductance (gs) and transpiration rate (E).Different water supplies greatlyinfluenced the tissue N concentration than P and K concentration.P.tricuspidata had a low Kconcentration (below 10 g.kg~(-1)) in all its plant tissues,which may allow the plants to limit itsstomatal opening and reduce its water loss.Water supplies had significantly affected N-useefficiency,but did not affect P and K-use efficiency.
9.P.tricuspidata had higher survival and growth rates,and showed well adaptability todisadvantageous growing conditions of rock slope so that it could be used as pioneer plant speciesto revegetation of rock slopes.The compress substrates can impove the biomass and growth rateof P.tricuspidata planting in rock slope.
引文
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