东南景天Sedum alfredii修复重金属污染土壤的研究进展

植物修复是净化重金属污染土壤最有潜力的原位修复技术之一,因地制宜地选择植物修复材料是大田植物修复成功实施的关键。国内外已发现的众多重金属超积累植物具有不同的生理特性,其中东南景天Sedum alfredii可忍耐和富集镉(Cd)和锌(Zn)等多种重金属,并且具有生长迅速的特点,是研究植物修复技术的模式植物之一。综述了东南景天对重金属的耐性、吸收、转运及解毒机制,阐明东南景天研究过程中存在的问题,并对其研究前景和方向作了简要分析,为重金属污染土壤植物修复工作的深入开展提供参考。

Growth Response and Metal Accumulation of Sedum alfredii to Cd/Zn Complex-Polluted Ion Levels

Sedum alfredii Hance has been identified as a new Zn-hyperaccumulator native to China. In this study, responses and metal accumulation of S alfredii were examined under Zn/Cd complex polluted conditions. The results showed that optimal growth of S alfredii in terms of the maximum dry matter yield was observed at Zn/Cd complex level of 500/100 mumol/L. Plant cadmium (Cd) or zinc (Zn) concentrations increased with increasing Cd or Zn supply. During the 20 d treatment, the highest Cd concentration in the leaves reached 12.1 g/kg at Zn/Cd level of 50/400 mumol/L and that of Zn in the stems was 23.2 g/kg at Zn/Cd level of 1000/50 mumol/L. The distribution of Cd in different plant parts decreased in the order: leaf > stem greater than or equal to root, whereas that of Zn was: stem > leaf greater than or equal to root. The accumulation of Cd and Zn in the shoots and roots of S. alfredii increased with the increasing of Zn/Cd supply levels, peaked at Zn/Cd levels of 250/400 and 500/100 mumol/L, respectively. The highest Cd and Zn uptake by the shoots was approximately 5 and 11 mg/plant, and was over 20 and 10 times higher than those in the roots, respectively. Zn supply at levels less than or equal to 500 mumol/L increased plant Cd concentrations, whereas high Zn supply decreased root Cd but did not affect leaf Cd concentrations in S alfredii Low Cd supply increased Zn concentration in the leaves, but Cd supply higher than 50 mumol/L considerably reduced root Zn concentrations, especially at low Zn level. These results indicate that S. alfredii can tolerate high Zn/Cd complex levels and has an extraordinary ability to hyperaccumulate not only Zn but also Cd. It could provide a new valuable plant material for understanding the mechanisms responsible for co-hyperaccumulation of Zn and Cd as well as for phytoremediation of the Cd/Zn complex polluted soils.

Sedum alfredii : A New Lead-Accumulating Ecotype

hi a survey of plant population, Sedum alfredii Hance, a new lead (Pb)-tolerant and lead (Pb)accumulating ecotype, was found in an old Pb/Zn mining area in Zhejiang Province of China. The growth and Pb content of plant ecotypes being able to and unable to accumulate Pb were studied by hydroponic culture with different concentrations of Pb(NO3)(2). Growth of shoots of accumulating ecotype was not affected by Ph treatments up to 320 mg/L, whereas that of non-accumulating ecotype was inhibited in all Ph treatments. The Ph concentrations in the roots and shoots of accumulating ecotype increased with increasing of Pb level in the nutrient solution. The maximum Ph concentrations in the shoots and roots of accumulating ecotype were 514 mg/kg and 13 922 mg/kg, 2.27 times and 2.62 times as much as that of non-accumulating ecotype, respectively. The highest rate of Pb accumulation of accumulating ecotype was 8.62 mug/plant/d, 7.16 times as much as that of non-accumulating ecotype. Due to its fast growth rate and high Pb-accumulating ability, from a phytoremediation perspective, accumulating ecotype of S. alfredii is a potential plant species for Pb removal from contaminated soils.

Subcellular Distribution of Cadmium in Mining Ecotype Sedum alfredii

The mining ecotype Sedum alfredii Hance could tolerate and grow normally in a nutritive solution containing cadmium (Cd) as high as 400 mumol/L. Under such a high Cd concentration, the subcellular accumulation of Cd in root, stem and leaf of this plant was found to be the highest in the cell wall, less in the soluble fraction and lowest in the cell organs. The mode of subcellular distribution of Cd in the mining ecotype S. alfredii was similar to other hyper accumulators of heavy metals, in which Cd was distributed more in the aerial part of plant. The results suggest that the mining ecotype S. alfredii is a new species of Cd hyperaccumulator.

两种生态型东南景天(Sedum alfredii Hance)根系互作效应对植株生长及吸收利用土壤Zn的影响

采用盆栽套种方法探讨了不同程度根系互作对两种生态型东南景天的生长和吸收利用土壤Zn的影响,以及超积累生态型东南景天的根系是否能活化土壤中难利用态Zn。不同程度的根系互作采用不同隔膜处理,即无隔膜处理、400目尼龙筛布隔开和PVC板隔开。结果发现,当两种生态型东南景天套种时,不同程度的根系互作对非超积累生态型东南景天的地上部和根系的生物量、根系Zn含量有显著影响。当与超积累生态型东南景天的根系完全交互作用或采用400目尼龙筛布隔开时,非超积累生态型的根系Zn含量显著增加,其地上部和根系生物量比其单种时显著减少。这些结果间接表明,超积累生态型东南景天根系能够活化土壤中的难溶态Zn。

Zinc adsorption and desorption characteristics in root cell wall involving zinc hyperaccumulation in Sedum alfredii Hance

Radiotracer techniques were employed to characterize 65Zn adsorption and desorption in root-cell-wall of hyperac-cumulating ecotype (HE) and non-hyperaccumulating ecotype (NHE) species of Sedum alfredii Hance. The results indicated that at the end of a 30 min short time radioisotope loading period, comparable amounts of 65Zn were accumulated in the roots of the two ecotypes Sedum alfredii, whereas 2.1-fold more 65Zn remains in NHE root after 45-min desorption. At the end of 60 min uptake period, no difference of 65Zn accumulation was observed in undesorbed root-cell-wall of Sedum alfredii. However, 3.0-fold more 65Zn accumulated in desorbed root-cell-wall of NHE. Zn2+ binding in root-cell-wall preparations of NHE was greater than that in HE under high Zn2+ concentration. All these results suggested that root-cell-wall of the two ecotypes Sedum alfredii had the same ability to adsorb Zn2+, whereas the desorption characteristics were different, and with most of 65Zn binding on root of HE being available for loading into the xylem, as a result, more 65Zn was translocated to the shoot.

Zinc removal from hyperaccumulator Sedum alfredii Hance biomass

Leaching of heavy metals from Sedum alfredii Hance biomass was studied with ammonia-ammonium chloride solution as leaching agent.The research was carried out in two phases:1)a leaching study to determine the zinc extraction efficiency of this leaching agent,and 2)a thermodynamic analysis to identify the likely reactions and stable Zn(Ⅱ)species formed in the leaching systems.Taguchi orthogonal experiment,with four variable parameters,leaching temperature,molar ratio of NH4Cl to NH3,leaching time and solid-to-liquid(L/S)ratio,and each at three levels,was used to optimize the experiment parameters by the analysis of variances.The results indicate that leaching temperature has the most dominant effect on metal extraction performance,followed by molar ratio of NH4Cl to NH3,solid-to-liquid ratio and leaching time.The optimum condition was obtained as follows:temperature of 60℃,molecular ratio of NH4Cl to NH3 of 0.6,leaching time of 2 h and solid-to-liquid ratio of 5-1.The total zinc leaching efficiency under optimum conditions reaches 97.95%.The thermodynamic study indicates that the dominant species produced by the leaching process should be the soluble Zn(NH3)4 2+.

Differential generation of hydrogen peroxide upon exposure to zinc and cadmium in the hyperaccumulating plant specie(Sedum alfredii Hance)

Sedum alfredii Hance has been identified as zinc(Zn) and cadmium(Cd) co-hyperaccumulator.In this paper the relationships of Zn or Cd hyperaccumulation to the generation and the role of H2O2 in Sedum alfredii H.were examined.The results show that Zn and Cd contents in the shoots of Sedum alfredii H.treated with 1000 μmol/L Zn2+ and/or 200 μmol/L Cd2+ increased linearly within 15 d.Contents of total S,glutathione(GSH) and H2O2 in shoots also increased within 15 d,and then decreased.Total S and GSH contents in shoots were higher under Cd2+ treatment than under Zn2+ treatment.However,reverse trends of H2O2 content in shoots were obtained,in which much higher H2O2 content was observed in Zn2+-treated shoots than in Cd2+-treated shoots.Similarly,the microscopic imaging of H2O2 accumulation in leaves using H2O2 probe technique showed that much higher H2O2 accumulation was observed in the Zn2+-treated leaf than in the Cd2+-treated one.These results suggest that there are different responses in the generation of H2O2 upon exposure to Zn2+ and Cd2+ for the hyperaccumulator Sedum alfredii H.And this is the first report that the generation of H2O2 may play an important role in Zn hyperaccumulation in the leaves.Our results also imply that GSH may play an important role in the detoxification of dissociated Zn/Cd and the generation of H2O2.

Integrated Fertilization Regimes Boost Heavy Metals Accumulation and Biomass of Sedum alfredii Hance

The hyperaccumulator Sedum alfredii Hance(S.alfredii)may be employed for zinc(Zn)and cadmium(Cd)-polluted soil remediation.However,the low phytoremediation efficiency,related to the low biomass production,limits its use with that purpose.In this experiment,nitrogen(N),phosphorus(P),and potassium(K)fertilizers,and organic manure were applied to investigate the phytoremediation ability of S.alfredii.Hydroponic and pot experiments were conducted using Zn-Cd polluted soil.The hydroponic experiment indicated that appropriate fertilizer application could increase(p<0.05)the amount of accumulated Zn and Cd in S.alfredii.When N supply ranged from 0.5 to 2.5 mmol L^(−1),it could improve growth and accumulation of Zn and Cd in whole plants of S.alfredii.The 1 mmol L^(−1) N was an optimal N dosage for shoot biomass production and Cd accumulation in shoots,while the 2.5 mmol L^(−1) was an optimal N dosage for Zn accumulation in shoots.Both low(<0.05 mmol L^(−1))and high(>0.8 mmol L^(−1))P supply decreased growth,and Zn/Cd accumulation in whole plants of the studied species.The 0.1 mmol L^(−1) P was an optimal dosage for S.alfredii biomass production and Zn/Cd accumulation in shoots.The supply levels within the range from 0.3 to 1 mmol L^(−1) K could significantly improve the biomass production of S.alfredii and its capability to accumulate Zn and Cd in the biomass.The 0.5 mmol L^(−1) K was an optimal dosage for the whole biomass production and Zn accumulation in shoots,while the 1 mmol L^(−1) was an optimal K dosage for Zn accumulation in shoots,which was 17.2%higher than the control.Moreover,the soil pot experiment showed that the combination of organic(fermented manure)and inorganic fertilizers made significant effects on the Zn and Cd-polluted soil remediation by S.alfredii.These effects varied,however,with the application of different proportions of N,P,K and organic matter.The Zn accumulation by S.alfredii reached the highest efficiency ability under the highest fertilizer mixing rate(N:50 mg kg^(−1),P:40 mg kg^(−1),K:100 mg kg^(−1),organic matter:1%).Even more,S.alfredii showed the strongest ability to accumulate Cd with a lower fertilizer mixing rate(N:25mg kg^(−1),P:20mg kg^(−1),K:50 mg kg^(−1),organic matter:0.5%).

Reactive oxygen species metabolism during the cadmium hyperaccumulation of a new hyperaccumulator Sedum alfredii (Crassulaceae)

Sedum alfredii Hance, a newly discovered hyperaccumulator, could serve as a good material for phytoremediation of Cd polluted sites. Malondialdehyde （MDA）, reactive oxygen species （ROS） and antioxidases （catalase （CAT）; superoxide dismutase （SOD）; peroxidase （POD）） in the leaf were determined when S. alfredii was treated for 15 d with various CdC12 concentrations ranging from 0 to 800 μmol/L. The results showed that the production rate of 2＇,7＇-dichlorofluorescein （DCF）, which is an indicator of ROS level, reached up to the maximum at 400 μmol/L CdCl2 and then declined with the increase of CdCl2 concentration, while MDA accumulation tended to increase. CAT activity was significantly inhibited at all tested CdCl2 concentrations and SOD activity was sharply suppressed at 800 μmol/L CdCl2. However, the enhancement of POD activity was observed when CdCl2 concentration was higher than 400 μmol/L. In addition, its activity increased when treated with 600 μmol/L CdCl2 for more than 5 d. When sodium benzoate, a free radical scavenger, was added, S. alfredii was a little more sensitive to Cd toxicity than that exposed to Cd alone, and the Cd accumulation tended to decline with the increase of sodium benzoate concentration. It came to the conclusions that POD played an important role during Cd hyperaccumulation, and the accumulation of ROS induced by Cd treatment might be involved in Cd hyperaccumulation.

Organic Materials Could Improve the Phytoremediation Efficiency of Soil Potentially Hazardous Metal by Sedum alfredii Hance

Soil potentially hazardous metal(PHM)is continually attracting public attention worldwide,due to its highly toxic properties and potentially huge damage to human being through food chain.Phytoremediation is an effective and eco-friendly way in remediation technology.A pot experiment was carried out to investigate the effect of different organic materials(biogas residue(BR),mushroom residue(MR),and bamboo-shoot shell(BS))application on phytoremediation of two PHM-contaminated soils(Fuyang soil as‘heavily-polluted soil’and Wenzhou soil as‘moderately-polluted soil’,respectively)by Sedum alfrecdii Hance.The results indicated:1)for moderately-polluted soil,the 5%BR treatment had the strongest activation to Cu and Zn,for heavily-polluted soil,1%BS treatment had the highest activation effect for Cu,Zn,Pb and Cd.2)the above-ground biomass of Sedum alfredii Hance increased with the addition rate of organic materials.3)for Cd uptake of Sedum alfredii Hance in moderately-polluted soil,only 1%BS treatment had a better accumulation effect,compared to the control,for Zn element,MR treatments were weaker than the control,while other treatments were better than the control,of which 5%BR,1%BS and 5%BS accumulated more Zn element by 39.6%,32.6%and 23.8%,respectively;in heavily-polluted soil,the treatments of 5%BS,1%BR and 5%BR accumulated more Cd than the control by 12.9%,12.8%and 6.2%,respectively,the treatments with organic materials addition promoted Zn accumulation in shoots of Sedum alfredii Hance,and the best treatment was 5%BS.Therefore,an appropriate application rate of BS and BR could improve the remediation efficiency for Zn/Cd contaminated soils by Sedum alfredii Hance.

植物套作系统修复镉污染农田土壤的效应——以苏南地区为例

为了研究不同套作模式对苏南地区Cd污染农田土壤的修复效果,选用能在苏南地区生长的伴矿景天、东南景天、苋菜和龙葵4种Cd超富集植物与大豆、油菜和小麦3种常规作物在Cd污染农田土壤中进行单作和套作修复试验.结果显示,与油菜套作后,4种超富集植物的Cd吸收量均有所增加,其中东南景天增幅最大,地上部Cd含量比单作增加24.4%;与小麦套作后,东南景天对Cd的吸收有所增加,地上部Cd含量比单作增加2.75%,龙葵和苋菜对Cd的吸收变化不显著,说明与油菜和小麦套作均能促进东南景天对Cd的吸收.套作伴矿景天减少了大豆根部Cd含量,套作苋菜和龙葵显著降低大豆茎、叶部分的Cd含量;套作模式下油菜根、茎的Cd含量均显著降低,叶片Cd含量受套作影响不显著;套作均显著降低了小麦的Cd含量,表明套作能抑制作物对Cd的吸收.此外,东南景天-油菜模式下Cd总积累量和土壤Cd去除率最高,为1343μg/盆和16.4%;东南景天-小麦次之,为1053μg/盆和9.8%.因此,东南景天-油菜/小麦两种套作模式均是苏南地区Cd污染农田土壤的适宜植物套作修复模式.研究结果可为该地区Cd污染农田修复同时安全利用提供科学依据和技术支持.

硫铅处理下伴矿景天对铅的耐受性与积累性研究

【目的】探明土壤在硫(S)铅(Pb)处理下,伴矿景天生长性状变化情况及对重金属Pb的耐受性与积累性,为Pb污染土壤植物修复提供依据。【方法】以伴矿景天为研究对象,采用随机区组试验,研究不同Pb浓度[Pb(CH_(3)COO)_(2)为0 mg、200 mg、400 mg、800 mg、1600 mg]和S浓度(Na_(2)SO_(4)为0 mg、50 mg、100 mg)对伴矿景天的株高、叶长、叶宽、根长、地上干重、根干重和Pb含量的影响及Pb耐受性和富集性的响应。【结果】在相同Pb浓度处理下,随S浓度增加,伴矿景天株高、叶长和根长整体呈增高趋势,在Pb 0 mg/kg、S 100 mg/kg处理下,根长最长,达6.50 cm。地上部干重以Pb 1600 mg/kg、S 50 mg/kg处理最小,根干重以Pb 1600 mg/kg、S 100 mg/kg处理最小。地上部Pb含量以Pb 1600 mg/kg、S 50 mg/kg处理最大,为1654.88 mg/kg;根系Pb含量以Pb 800 mg/kg、S 50 mg/kg处理最大,为1070.42 mg/kg。伴矿景天地上部干重与根干重呈极显著正相关关系,叶长与叶宽呈极显著正相关关系。伴矿景天的Pb富集系数在相同S浓度下,随Pb浓度增加而减小,转运系数、转运量系数总体随Pb浓度增加而减小;在相同Pb浓度处理下,富集系数随S浓度增高而增加,富集系数、转运系数和转运量系数、迁移率最高分别为11.04、2.88、30.64和74.24%。【结论】伴矿景天在铅胁迫下外施硫可缓解地上部、根生物量和株高的减少,并提高植株对铅的富集系数,外施硫是促进伴矿景天吸收重金属的有力措施。

菌肥及泥炭土浸提液对植物提取土壤Cd的影响

采用温室盆栽试验,研究菌肥和泥炭土浸提液对东南景天生物量、Cd富集特征、土壤Cd有效性和土壤理化性质及对后茬水稻在落干条件下水稻Cd的影响。结果表明,菌肥和泥炭土浸提液均能提高东南景天对土壤重金属的提取效率,其中施用两次液体菌肥的效果最好,东南景天对Cd提取量为0.34 mg·pot^(-1),是对照的3.1倍,泥炭土浸提液处理的东南景天Cd提取量为0.32 mg·pot^(-1),是对照的2.9倍,这两种处理对Cd的提取率较高,达到13%~15%,土壤全量Cd的降低幅度达到17.55%~20.41%。东南景天收获后种植水稻,落干条件下促进了水稻对Cd的吸收,稻米和茎叶的Cd提取量较对照增幅分别为30.31%~396.24%和12.36%~257.65%,其中在菌肥及泥炭土浸提液联合处理下水稻对Cd的吸收最大,稻米和茎叶的Cd含量分别为2.14 mg·kg^(-1)和2.88 mg·kg^(-1),水稻地上部对Cd总提取率达到3.15%。因此,施用两次菌肥是提高东南景天提取土壤Cd的有效措施,菌肥与泥炭土浸提液联合施用在落干条件下显著强化了水稻对土壤Cd的提取。

Differences of Uptake and Accumulation of Zinc in Four Species of Sedum

Four species of Sedum L. were grown in nutrient solution for the comparison of their Zn uptake and accumulation. S. alfredii Hance showed much greater tolerance to Zn than the other three species. Shoot and root yields of S. sarmentosum Bunge, S. bulbiferum Makino, and S. emarginatum Migo decreased with increasing Zn concentration in the solution, while shoot and root yields of S. alfredii increased when Zn concentration was ≤80 mg·L -1. At 80 mg·L -1 Zn, Zn concentration in shoots of S. alfredii reached 19.09 mg·g -1. S. alfredii was also more efficient in Zn translocation from roots to shoots, while Zn concentration in shoots was much higher than in roots. However, this was not the case for the other three species. The results showed that S. alfredii is a Zn hyperaccumulator and could be useful for the phytoremediation of Zn-contaminated soils.

不同预处理和干燥方式下养心菜粉色泽、功能成分和抗氧化性水平的比较

为了研究养心菜的加工适用性,拓展其利用途径,采用未预处理(Untreated,UT)、沸水漂烫(Boiling Water Blanching,BWB)、沸水+护色漂烫(COLOR Protection+Water Blanching,CPB)、蒸汽漂烫(Steam Blanching,SB)与冻融(Freeze-Thawing,FT)五种预处理方式处理后,分别进行热风干燥(Hot Air Drying,HA)、热泵干燥(Heat Pump Drying,HP),得到UT HP、UT HA、BWB HP、BWB HA、CPB HP、CPB HA、SB HP、SB HA、FT HP及FT HA 10种养心菜粉,比较不同预处理方式和干燥方式下养心菜色泽、功能成分及抗氧化能力水平的变化。结果表明:UT HP没食子酸和山萘酚含量最高,但色泽差、叶绿素含量较低,干燥时间长;CPB HP的干燥时间短,Vc含量最高,具有较高的抗氧化水平,但没食子酸含量较低;SB HP总酚、总黄酮、槲皮素含量最高,其他品质指标介于中间;FT HP养心菜具有最好的色泽、最高的叶绿素、游离氨基酸含量,但总酚、总黄酮、槲皮素、山萘酚以及抗氧化水平低。从主成分分析来看,BWB HP、CPB HP、SB HP和BWB HA、CPB HA、SB HA的养心菜品质分别具有一定的相似性,UT HP、FT HP、UT HA、UT HA和其有明显差异。该文综合比较了养心菜不同预处理及干燥方法的优劣,在养心菜下游的功能性食品的开发过程中,可以根据需求选择合适的干燥方法。

菜心与东南景天间作修复镉污染农田的生态效应

为了探究镉低积累型菜心碧绿粗苔和镉超积累植物东南景天间作修复镉污染土壤的效果,在华南农业大学生态农场进行了模拟控制试验,分别采集菜心、东南景天在单作和间作时的根际土进行研究。结果表明:间作可显著提高植物的生物量,其中东南景天地上和地下部的生物量较单作分别提高了79.05%和50.00%,菜心地上和地下部的生物量则分别提高了110.96%和125.00%;间作使东南景天地上和地下部的镉含量较单作分别提高了7.61%和70.73%,但菜心地上部的镉含量较单作降低了27.21%;东南景天茎叶储存Cd的主要亚细胞结构为液泡中的细胞液,占比高达90%,远高于细胞器与细胞壁中的Cd含量;间作显著增加了土壤的pH,相较于东南景天、菜心单作时的pH分别增加了0.39、0.28个单位;间作降低了菜心根际土镉的有效性,较单作低19.23%;另外,间作可使土壤微生物总量和蔗糖酶活性显著增加,其中放线菌数量比菜心、东南景天单作时分别增加了27.95%、24.28%,蔗糖酶活性则分别提高了164.89%、88.48%。同时,通过计算可知间作的东南景天土壤镉提取率是单作的1.88倍。综上所述,镉低积累型菜心与镉超积累植物东南景天间作,不仅可提高菜心生产的安全性,而且增加了东南景天对土壤镉的提取效率,是一种经济、有效且风险较低的边生产边修复的模式。

江西景天属一新种——溪水景天

【目的】2021年在中国江西东北部阳际峰国家级自然保护区进行林木种质资源调查时发现一种景天科(Crassulaceae)景天属(Sedum L.)植物,该种与已报道的同属植物均有区别,本文予以论述并确定其分类学地位。【方法】跟踪阳际峰保护区多个种群,在不同物候期进行详细观察和形态学比较,确定其相近种及区别特征,进而通过文献检索和标本查询确定其在全国分布情况。【结果】该种营养枝在幼苗阶段呈莲座状,最为相似种为细小景天(Sedum subtile Miq.),区别特征为新发现种具有发达且3叶轮生的不育枝、花茎上每分枝5~16朵花,苞片匙状倒披针形;细小景天不育枝细弱且叶互生或对生、花茎每分枝3~5朵花、苞片线形。【结论】该种为景天属一新种,命名为“溪水景天”(Sedum fluviale B.Chen&Z.W.Zhu)。该种专一性地生于溪流环境,分布范围较广,包括江西、浙江、福建、安徽、湖南、贵州等省(区、市),此前存在较多错误鉴定的情况。模式标本存放于上海辰山植物标本馆(CSH)。

景天属(Sedum)8种植物茎的解剖学研究

利用光学显微镜 ,首次观察并报道了安徽产景天属 (Sedum ) 8种植物茎的解剖结构 ;结果表明 ,景天属植物茎由表皮 (一层 )、皮层 (数层 )、维管柱 (木质部、韧皮部、髓 )三部分构成 ,并明确提出了肉质植物茎的解剖学处理方法 .