都市及工業廢水常會藉由灌溉渠道進入農田，造成農田汙染，重金屬的議題被關心主因為重金屬無法被自然中生物所分解以及對人體健康的潛在影響，在台灣許多工廠排水系統為灌排不分離，導致廢水經由灌溉渠道排入農田造成如Cd、Zn、Cu、Cr、Ni及Pb等重金屬造成土壤污染，當廢水中含有重金屬進入土體，造成地下水污染或藉由土壤溶液被植物所吸收。本研究模以灌溉水pH值的改變對土壤性質與水稻吸收重金屬(Cd、Zn、Cu、Cr、Ni、Pb)之影響，研究中使用來自受工業區廢水污染之土壤，以盆栽試驗探討pH值受重金屬污染土壤之影響，結果顯示酸性灌溉水對於稻米成長影響甚小，不論地上部或地下部之植體，所測得之重金屬鎘、鉻、銅及鋅呈現差異顯著，且當pH 值調降愈低時，植體中重金屬濃度愈高，故調降灌溉水pH 值會影響植體中重金屬含量。從土壤中重金屬濃度發現鉻在盆栽試驗中皆呈顯著差異，重金屬中最容易受pH 值之影響。98 年二期作土壤鎘在pH 值調降3 隔次處理時濃度最低，但在糙米及莖葉鎘在pH 值調降3 隔次處理時濃度最高，顯示出此處理造成土壤中鎘溶出最多進而被植體吸收造成此現象。98 年二期作中探討植體與土壤重金屬濃度與低分子量有機酸之相關性，結果顯示糙米鉻濃度提高時，草酸及乙醛酸濃度隨之增加，鋅濃度增加時，草酸濃度也增加；而莖葉鎘濃度提高時，反丁烯二酸及蘋果酸濃度也隨之增加另外根部鉛及鉻濃度增加時，乙醛酸濃度也隨之增加且當土壤中銅濃度較高時，左旋乳酸也隨之增加。

The use of industrial or municipal wastewater to the farm land by irrigation channels causes the pollution of cultivated agricultural lands. Among various pollutants heavy metals are the most serious concerned because of their nonbiodegradable nature and potential impact on human heath. Many factories in Taiwan discharged their effluents/wastewater into drainage and supplied to the farm lands through irrigation channels, since there is no separate system for drainage and irrigation channels. It is known that soils irrigated by wastewater accumulate heavy metals such as Cd, Zn, Cu, Cr, Ni and Pb in surface soil. When the capacity of the soil to retain heavy metals is reduced due to repeated use of wastewater, soil can release heavy metals into ground water or soil solution available for plant uptake. This study describes the effect of pH changes of irrigation water on soil heavy metal pollution from industrial and municipal wastewater and subsequently the uptake of Cd, Zn, Cu, Cr, Ni and Pb heavy metals by rice plant have been investigated. In this study, soil was collected from polluted farm land which is irrigated with wastewater from industrial and urban areas. Pot experiments were conducted by growing rice plants to study the effect of pH on soil heavy metal pollution at original irrigation water pH (around at 7.0) and below original irrigation water pH (1.5 and 3.0) and found that Cd, Cr, Cu and Zn exhibit significant difference with the change of pH values. When the pH value of irrigation water is lower than original irrigation water pH, the observed concentration of heavy metals uptake by all parts of rice plant was higher. After crop production was over, the stems and leaves of the rice plants and soil were examined to determine the concentrations of heavy metals and found that cadmium was highest concentration among studied heavy metals at pH value lower than original irrigation water pH compared to other pH values. It indicates that rice plant highly uptake of cadmium from contaminated soil which is irrigated by wastewater. The results showed that the soil was affected by acidic irrigation water and uptake of the heavy metals by rice plant significantly different with the pH values. During the growth of rice plant, effect of low molecular weight organic acids on heavy metal pollution in different parts of rice plant was determined. The results showed that Cr concentration was higher in rice, the oxalic acid and glyoxylic acid concentration in soil was also higher observed. In stems of the rice plant the concentration of Cd positively correlated with the concentrations of fumaric and malic acid. Glyoxylic acid concentration in soil was correlated with the concentrations of Pb and Cr in roots of rice plant. In soil the concentration of Cu correlated with L-(+) Lactic acid concentration.

目 錄
摘 要 Ⅰ
Abstract Ⅲ
誌 謝 V
目 錄 VI
表 目 錄 X
圖 目 錄 XI
第一章 前言 1
1.1 研究緣起 1
1.2 研究目的 3
第二章 文獻回顧 5
2.1 重金屬污染對環境的影響 5
2.1.1重金屬污染來源 5
2.1.2重金屬之特性與危害 10
2.2影響土壤中重金屬之傳輸因子 17
2.3土壤中重金屬之固相鍵結型態及其分佈 19
2.4根圈低分子量有機酸之特性與定量 22
2.4.1根圈土壤(rhizosphere soil)的定義與特性 22
2.4.2有機酸之特性 23
2.4.3有機酸的分析 24
2.4.4低分子量有機酸含量測定 25
2.4.5低分子量有機酸與重金屬之關係 26
第三章 材料與方法 27
3.1實驗設備材料 27
3.1.1供試土壤 27
3.1.2儀器與藥品 27
3.2試驗方法 27
3.2.1土壤及灌溉水體基本性質分析方法 27
3.2.2植體中重金屬含量分析方法 33
3.3根圈土壤溶液LMWOAs之萃取 33
3.3.1根圈土壤溶液之揮發性LMWOAs 萃取 33
3.3.2 LMWOAs 標準品 33
3.4 LMWOAs分析之氣相層析儀之規格與設定 33
3.4.1揮發性LMWOAs之分析 33
3.4.2非揮發性LMWOAs之衍生化 33
3.4.3非揮發性LMWOAs之分析 34
3.4.4 LMWOAs滯留時間(retention time)之確立 34

第四章 結果與討論 41
4.1 調降灌溉水pH 值對稻米生長之影響 41
4.2 97 年二期作盆栽試驗 45
4.2.1 97 年二期作水稻糙米重金屬含量 45
4.2.2 97 年二期作水稻莖葉重金屬含量 46
4.2.3 97 年二期作水稻根部重金屬含量 48
4.2.4 97 年二期作土壤重金屬含量 49
4.2.5 97 年二期作土壤0.1 M 鹽酸萃取量 51
4.3 98 年一期作盆栽試驗 53
4.3.1 98 年一期作水稻糙米重金屬含量 53
4.3.2 98 年一期作水稻莖葉重金屬含量 54
4.3.3 98 年一期作水稻根部重金屬含量 55
4.3.4 98 年一期作水稻土壤重金屬含量 56
4.3.5 98 年一期作土壤0.1 M 鹽酸萃取量 57
4.4 98 年二期作盆栽試驗 58
4.4.1 98 年二期作水稻糙米重金屬含量 58
4.4.2 98 年二期作水稻莖葉重金屬含量 59
4.4.3 98 年二期作水稻根部重金屬含量 60
4.4.4 98 年二期作土壤重金屬含量 60
4.4.5 98 年二期作土壤0.1 M 鹽酸萃取量 62
4.5 98 年二期作水稻重金屬含量與非揮發性有機酸之變化 63
4.5.1 98 年二期作水稻糙米重金屬含量與非揮發性有機酸之變化
63
4.5.2 98 年二期作水稻莖葉重金屬含量與非揮發性有機酸之變化
68
4.5.3 98 年二期作水稻根部重金屬含量與非揮發性有機酸之變化
71
4.5.4 98 年二期作盆栽土壤重金屬含量與非揮發性有機酸之變化
76
第五章 結論 81
參考文獻 82
附錄 88


表目錄
表2-1 土壤重金屬污染監測基準 6
表2-2 土壤重金屬污染管制標準 7
表2-3 國內土壤中重金屬主要來源說明表 9
表2-4 重金屬在土壤及植物中的臨界毒性 10
表3-1供試土壤及灌溉水之重金屬基本特性分析 31
表3-2 供試土壤及灌溉水之基本特性分析 32
表3-3 研究中所使用之儀器廠牌與型號 35
表3-4 研究中所使用之藥品與廠牌 36
表3-5 測定之低分子量有機酸之揮發性質、種類、化學式及中文名
37
表3-6 測定之低分子量有機酸之滯留時間 38
表4-1 97 年二期作溫室盆栽稻作糙米之千粒重、株高與分糵數 42
表4-2 97 年二期作田間盆栽稻作糙米之千粒重、株高與分糵數 42
表4-3 98 年一期作溫室稻作糙米之千粒重、株高及分糵數 43
表4-4 98 年二期作溫室稻作糙米之千粒重、株高及分糵數 44


圖目錄
圖1-1 研究架構圖 4
圖3-1 非揮發性有機酸之氣相層析分析滯留時間 39
圖3-2 揮發性有機酸之氣相層析分析滯留時間 40
圖4-1 97 年二期作溫室水稻糙米重金屬含量 45
圖4-2 97年二期作田間水稻糙米金屬含量 46
圖4-3 97 年二期作溫室水稻莖葉重金屬含量 47
圖4-4 97 年二期作田間水稻莖葉重金屬含量 47
圖4-5 97 年二期作溫室水稻根部重金屬含量 48
圖4-6 97 年二期作田間水稻根部重金屬含量 49
圖4-7 97 年二期作溫室盆栽土壤王水萃取量 50
圖4-8 97 年二期作田間盆栽土壤王水萃取量 50
圖4-9 97年二期作溫室盆栽土壤0.1 M 鹽酸萃取量 52
圖4-10 97 年二期作田間盆栽土壤0.1 M 鹽酸萃取量 52
圖4-11 98 年一期作溫室水稻糙米重金屬含量 53
圖4-12 98 年一期作溫室水稻莖葉重金屬含量 54
圖4-13 98 年一期作溫室水稻根部重金屬含量 55
圖4-14 98 年一期作溫室盆栽土壤王水萃取量 56
圖4-15 98 年一期作溫室盆栽土壤0.1 M 鹽酸萃取量 57
圖4-16 98年二期作溫室水稻糙米重金屬含量 58
圖4-17 98 年二期作溫室水稻莖葉重金屬含量 59
圖4-18 98 年二期作溫室水稻根部重金屬含量 60
圖4-19 98 年二期作溫室盆栽土壤王水萃取量 61
圖4-20 98 年二期作溫式盆栽土壤0.1 M 鹽酸萃取量 62
圖4-21 98 年二期作水稻糙米中鎘與左旋-乳酸、草酸及檸檬酸之比較 64
圖4-22 98 年二期作水稻糙米中鎘與羧基二酸、反丁烯二酸及乙醛酸之比較 64
圖4-23 98 年二期作水稻糙米中鎘與丙二酸、順丁烯二酸及草醋酸之比較 65
圖4-24 98 年二期作水稻糙米中鎘與蘋果酸、琥珀酸及右旋-酒石酸之比較 65
圖4-25 98 年二期作水稻糙米中鉻、銅、鎳及鋅與左旋-乳酸、草酸及右旋-檸檬酸之比較 66
圖4-26 98 年二期作水稻糙米中鉻、銅、鎳及鋅與羧基二酸、反丁烯二酸及乙醛酸之比較 66
圖4-27 98 年二期作水稻糙米中鉻、銅、鎳及鋅與丙二酸、順丁烯二酸及草醋酸之比較 67
圖4-28 98 年二期作水稻糙米中鉻、銅、鎳及鋅與蘋果酸、琥珀酸及右旋-酒石酸之比較 67
圖4-29 98 年二期作水稻莖葉中鎘及鉛與左旋-乳酸、草酸及檸檬酸之比較 68
圖4-30 98 年二期作水稻莖葉中鎘及鉛與羧基二酸、乙醛酸及反丁烯二酸之比較 69
圖4-31 98 年二期作水稻莖葉中鎘及鉛與羧基二酸、乙醛酸及反丁烯二酸之比較 69
圖4-32 98 年二期作水稻莖葉中鎘及鉛與琥珀酸、蘋果酸及右旋-酒石酸之比較 70
圖4-33 98 年二期作水稻莖葉中鉻、銅、鎳及鋅與左旋-乳酸、草酸及檸檬酸之比較 70
圖4-34 98 年二期作水稻莖葉中鉻、銅、鎳及鋅與羧基二酸、乙醛酸及反丁烯二酸之比較 71
圖4-35 98 年二期作水稻根部中鎘及鉛與左旋-乳酸、草酸及檸檬酸之比較 72
圖4-36 98 年二期作水稻根部中鎘及鉛與羧基二酸、反丁烯酸及乙醛酸之比較 72

圖4-37 98 年二期作水稻根部中鎘及鉛與丙二酸、順丁烯二酸及草醋酸之比較 73
圖4-38 98 年二期作水稻根部中鎘及鉛與蘋果酸、琥珀酸及右旋-酒石酸之比較 73
圖4-39 98 年二期作水稻根部中鉻、銅、鎳及鋅與左旋-乳酸、草酸及檸檬酸之比較 74
圖4-40 98 年二期作水稻根部中鉻、銅、鎳及鋅與羧基二酸、反丁烯二酸及乙醛酸之比較 74
圖4-41 98 年二期作水稻根部中鉻、銅、鎳及鋅與丙二酸、順丁烯二酸及草醋酸之比較 75
圖4-42 98 年二期作水稻根部中鉻、銅、鎳及鋅與蘋果酸、琥珀酸及右旋-酒石酸之比較 75
圖4-43 98 年二期作盆栽土壤中鎘與左旋-乳酸、草酸及檸檬酸之比較 77
圖4-44 98 年二期作盆栽土壤中鎘與羧基二酸、乙醛酸及反丁烯二酸之比較 77
圖4-45 98 年二期作盆栽土壤中鎘與丙二酸、順丁烯二酸及草醋酸之比較 78

圖4-46 98 年二期作盆栽土壤中鎘與琥珀酸、蘋果酸及右旋-酒石酸之比較 78
圖4-47 98 年二期作盆栽土壤中鉻、銅、鎳、鉛及鋅與左旋-乳酸、草酸及檸檬酸之比較 79
圖4-48 98 年二期作盆栽土壤中鉻、銅、鎳、鉛及鋅與羧基二酸、乙醛酸及反丁烯二酸之比較 79
圖4-49 98 年二期作盆栽土壤中鉻、銅、鎳、鉛及鋅與丙二酸、草醋酸及順丁烯二酸之比 80
圖4-50 98 年二期作盆栽土壤中鉻、銅、鎳、鉛及鋅與琥珀酸、蘋果酸及右旋-酒石酸之比較 80

參考文獻
于迺文、林正鈁，「土壤中重金屬含量的變化 (二) 土壤中重金屬的移除與
土地負荷量」，工業污染防治，第79期，第37-54頁 (2001b)。
王一雄、陳尊賢、李達源，「土壤污染學」，國立空中大學，台北，第30-35；
146-150頁 (1995)。
江漢全、鄒燦陽、張昭駿，「重金屬廠址土壤中鎳鉻之移動潛勢評估」，第
一屆土壤及地下水處理技術研討會 (2003)。
呂昱志，「重金屬污染農地之整治方案研擬-應用能值分析評估鎘污染農地整治方案」，碩士論文，國立中興大學資源管理研究所，台中 (1996)。
何勝惟，「應用多變量統計於土壤重金屬污染與土地利用之分析」，碩士論
文，國立台灣大學，台北 (2002)。
林宏達，「以植生復育技術試驗遭受溢油污染濕地土壤之研究」，碩士論文，
國立中山大學，高雄 (2004)。
翁震炘，「農作物重金屬污染監測與管制措施」，農政與農情，第169期，
第1-3頁 (2006)。
陳冠博，「受污染農地重金屬存在形態與環境因子之相關性研究」，碩士論
文，逢甲大學環境工程與科學研究所，第3-9頁 (2003)。
陳尊賢，「受重金屬污染土壤之整治技術與經驗」，土壤及地下水污染整治
技術研究論文集，第145-162頁 (2000)。
陳鴻堂，土壤與作物的鋅元素 (2004)。
郭魁士，土壤學，中國書局，台北市，第269-290；421-424頁 (1997)。
張如燕，「銅、鋅、鎘、鉛在受污染土體中之移動性及在土壤各組成分中之
分佈」，碩士論文，國立台灣大學，台北 (1989)。
張仁福，土壤污染防治學，復文書局，高雄市，第325-336頁 (1998)。
張添晉、郭文陽，「酸萃取、淋洗受重金屬污染土壤之研究」，第十五屆廢棄物處理技術研討會論文集 (2000)。
童翔新、陳冠博、黃舒瑜、劉忠裕、姜宇駿、陸運姵，「受污染農地重金屬
污染物之化學性質探討」，第二屆土壤及地下水處理技術研討會(2004)。
廖永綜，「蒸散作用對萵苣吸收鉛及根圈水溶性低分子量有機酸之影響」，
博士論文，國立中興大學土壤環境科學系，台中 (2006)。
劉黔蘭、王銀波，「鎳污染臺灣各類土壤與作物生長關係之研究」，第二屆
土壤污染防治研討會，第243-255頁 (1992)。
歐育憲，「土壤重金屬污染物之生物有效性意義研究」，碩士論文，逢甲大
學土木及水利工程研究所，台中，第4-13頁 (2000)。
蘇惠靖，「以萃取法復育受重金屬污染土壤之可行性評估」，屏東科技
大學環境工程與科學系，屏東 (2003)。

Alloway, B. J., “Heavy metal in soil, 2nd ed,” John Wiley & Sons, New
York (1995).
Alloway, B. J., “Soil processes and the behavior of heavy metals,”Heavy
metals in soil, John Wiley and Sons, New York, pp. 11-35 (1995).
Adhikari, T. and Singh, M. V., “Sorption characteristics of lead and cadmium
in some soil of India”, Geoderma, Vol. 114, pp. 81-92 (2003).
Broos, K., Uyttebroek, M., Mertens, J. and Somolders, E., “A survey of
symbiotic nitrogen fixation by white clover grown on matal contaminatsd
soil”, Soil Biology & Biochemistry, Vol. 36, pp. 633-640 (2004).
Brett, C. M. A. and Brett A. M. O., “Electrochemistry principle methods and
application”, Oxford University Press Inc., United Statres (1993).
Bradl, H. B., “Adsorption of heavy metal ions on soil and soils constituents”,
Journal of Colloid and Interface Science, Vol. 277, pp. 1-18 (2004).
Cushumsn, J. H.,“Nutrient transport insid and outside the
rhizosphere:Theory,”Soil Sci. Soc. Am. J., Vol. 46, pp. 704-709 (1982).
Chen, M. and Ma, L. O. “Comparison of Three Aqua Regia Digestion Methods
for Twenty Florida Soil.”Soil Sci. J. 65, pp. 491-499 (2001).
Chlopecka, A., “Assessment of form of Cd, Zn and Pb in contaminated
calcareous and gleyed soil in Southwest Poland,”Sci. Total Environ., Vol.
188, pp. 253-262 (1996).
Chaney, R. L., and Giordano, P. M., “Soils for management of organic wastes
and wastewaters,” Microelements as related to plant deficiencies and
toxicities,In L. F. Elliot and F. J. Stevenson (ed.) ASA, Madison, WI.,
pp.234-279 (1997).
Delhaize, E., Craig, S., Beaton, C. D., Bennet, R. J., Jagadish, V. C., and
Randall, P. J., “Aluminum tolerance in wheat (Triticum aestivum L.). I.
Uptake and distribution of aluminum in root apices,” Plant
Physiol.,Vol.103,pp 685-693 (1993).
Foy, C. D., Lee, E. H., Coradetti, C. C., and Taylor, G. J., “Plant
nutrition-physiology and applications,” Organic acids related to differential
aluminium tolerance in wheat (Triticum aestivum) cultivars,In M. L. van
Beusichem (ed.) Kluwer Academic Publishers, Boston., pp. 381-389(1990).
Fazeli, M.S., Sathyanarayan, Satish, P.N., Effect of paper mill effluents on
accumulation of heavy metals in coconut trees near Ninjangud, Mysore
District, Karnataks. India. J. Envi. Geo. Water Sci. 17 (1), 47–50 (1991).
Hammer, D., and Keller, C., “Changes in the rhizosphere of
metal-accumulating plant evidenced by chemical extractants,” J. Environ.
Qual., Vol. 31, PP.1561-1569 (2002).
Hantke, K., “Bacterial Zinc transporters and regulators,” BioMetals, Vol. 14,
pp.239-249 (2000).
Klassen, S. P., J. E. McLean, P. R. Grossl, and Sims, R. C., “Fate and behavior
of lead in soils planted with metal-resistant species (river birch and
smallwing sedge),” J. Environ. Qual., Vol. 29, pp. 1826-1834 (2000).
Krishanmurti, G. S. R., Cieoelinski, G., Huang, P. M., and Van Rees, K. C. J.,
“Kinetics of cadmium release form soil as influenced by organic acids:
Implication in cadmium availability,” J. Environ. Qual., Vol. 26, pp.
271-277 (1997).
Mcbride, M. B., “Environmental chemistry of soil,” Oxford University Press,
New York (1994).
Mench, M., Morel, J. L., Guckert, A., and Guillet, B., “Metal binding with root
exudates of low molecular weight,” J.Soil Sci., Vol. 39, pp.521-527(1988).
Naidu, R., and Harter, R. D., “Effect of different organic liands on cadmium
Sorption by and extractability from soil,” Soil Sci. Am. J., Vol. 62,
pp.644-650 (1998).
Nigam, R., Srivastave, S., Prakash, S., and Srivastava, M. M., “Cadmium
mobilization and plant availability-the impact of organic acids commonly
exuded from roots,” Plant Soil., Vol. 230, pp. 107-113 (2001).
Poulsen, I. F., and Hansen, H. C. B., “Soil sorption of nickel in presence of
citrate or arginine,” Water air soil pollut., Vol. 120, pp. 249-259 (2000).
Peter, R. W., “Chelant extraction of heavy metals from contaminated soils,” J.
Hazard. Mater., Vol. 66, pp. 151-210 (1999).
Salt, D. E., Blaylock, M., Kumer, N. P. B. A., Dushenkov, V., Ensley, B. D.,
Chet, I. and Raskin, I., “Phytoremediation: A novel strategy for the
removal of toxic metals form the environment using plant,”
Biotechnology, Vol. 13, pp. 468-474 (1995).
Szmigielska, A. M., Van Rees, K. C. J., Cieslinski, G., and Huang, P. M., “Low
Molecular weight dicarboxylic acids in rhizosphere soil of durum wheat,”
J. Agric. Food Chem., Vol.44, pp. 1036-1040 (1996).
Salt, D. E. Prince, R. C., Pickering, I. J., and Raskin, I., “Mechanisms of
cadmium mobility and accumulation in indian mustard,” Plant Physiol.,
Vol. 109, pp. 1427-1433 (1995).
Szmigielska, A. M., Van Rees, K. C. J., Cieslinski, G., and Huang, P. M.,
“Comparisom of liquid and gas chromatography for analysis of low
molecular weight organic acids in rhizosphere soil,” Commun. Soil Sci.
Plant Anal., Vol. 28, pp. 99-111 (1997).

Streit, B, Stumm, W. Chemical properties of metals and the process of bioaccumulation in terrestrial plants. Chapter 2 in Markert B. Plants as biomonitors, indicators for heavy metals in the terrestrial environment. Weinheim: VCH, pp. 2-31 (1993).
Tessier, A., P. G. C. Campbell and M. Bisson, “Sequential extraction procedure
for the speciation of particulate trace mtals,” Anal. Chem., Vol.51, pp.
844-855 (1979).
Taniguchi, S., Yamagata, N., and Sakurai, K., “Cadmium adsorption on
hydroxyaluminosilicate-montmorillonite complex as influenced by oxalate
and citrate,” Soil Sci. Plant Nutr., Vol. 46, pp. 315-324 (2000).
Uren, N. C. and Reisenauer, H. M., “The role of root exudates in nutrient
acquisition,” Adv. Plant Nutr., Vol. 3, pp. 79-114 (1988)
Veeken, A. and Hamelers, B., “Sources of Cd, Cu, Pb and Zn in biowaste,” The Science of the Total Environment., Vol. 300, pp.87-98 (2002).
Wu, Z. H., Gu, Z. M., Wang, X. R., Evans, L., Guo, H. Y., “Effect of organic
Acids on adsorption of lead onto montmorillonite, goethite and humic
acid,” Environ. Pollut. Vol. 121, pp. 469-475 (2003).
Wu, L.H., Luo, Y.M., Christie, P., Wong, M.H., “ Effects of EDTA and low
molecular weight organic acids on soil solution properties of a
heavy metal polluted soil,” Chemosphere , Vol. 50, pp. 819-822 (2003).
Welch, R. M., Micronutrient nutrition of plants. Crit Rev Plant
Sci14:49-82 (1995).
Zhao, F. J., Lombi, E., Breedon, T., and McGrath, S. P., “Zinc
Hyperaccumulation and cellular distribution in Arabidopsis halleri, “ Plant
Cell Environ., Vol. 23, pp. 507-514 (2000).