고축적식물의 중금속 흡수기작과 뿌리에 의한 근권 토양의 화학변화 - 총설 Hyperaccumulation mechanism in plants and the effects of roots on rhizosphere soil chemistry - A critical review원문보기
토양중 중금속을 흡수해서 체내에 고농도로 축적할 수 있는 식물, 이른바 고축적식물(hyperaccumulator)의 발견으로 오염토양에 대한 식물복원(phytoremediation) 기술에 대한 많은 연구들이 수행되고 있다. 이들 연구의 방향은 크게 고축적식물의 중금속 축적 기작을 밝히기 위한 것과 축적효율을 높임으로써 복원 효율을 향상시키는 실용적인 기술개발로 나누어진다. 지금까지 고축적식물에 의한 중금속 축적 기작은 다섯 가지의 특이 기작으로 알려져 있는데, 1) 뿌리세포의 중금속 흡수 증진, 2) 식물체 조직내의 중금속 이동성 향상, 3) 중금속의 무독화(detoxification) 및 격리(sequestration), 4) 토양-뿌리 경계면에서의 중금속 유효도 증진, 그리고 5) 중금속 오염토양으로의 능동적인 뿌리의 성장 등이 이에 속한다. 일반적으로 토양 중 낮은 중금속 유효도는 식물복원 기술의 현장 적용에 있어 제한요소로 간주된다. 이를 극복하기 위해서는 위에 기술된 다섯 가지 기작 중 고축적식물의 뿌리가 근권 토양중 중금속의 화학변화에 미치는 영향을 이해하는 것이 매우 중요하다. 식물 뿌리에 의한 근권 토양의 pH 변화와 뿌리에서 나오는 분자량이 적은 유기산(low-molecular-weight organic acids, LMWOAs)과 같은 유기성 분비물은 근권부 토양의 화학적 특성을 변화시키고 결과적으로 중금속의 유효도를 변화시킨다. 예를 들어 뿌리에서 나오는 $H^+$ 이온은 토양 pH를 감소시키고 이에 따라 중금속의 유효도는 증가한다. 또한 고농도의 중금속에 노출된 뿌리는 많은 양의 유기물질을 분비하게 되고 근권 토양에 축적되는 이 유기물질은 토양중 중금속과 결합하여 유기복합물질(organo-metallic complexes)을 형성하면서 유효도를 증가시킨다.
토양중 중금속을 흡수해서 체내에 고농도로 축적할 수 있는 식물, 이른바 고축적식물(hyperaccumulator)의 발견으로 오염토양에 대한 식물복원(phytoremediation) 기술에 대한 많은 연구들이 수행되고 있다. 이들 연구의 방향은 크게 고축적식물의 중금속 축적 기작을 밝히기 위한 것과 축적효율을 높임으로써 복원 효율을 향상시키는 실용적인 기술개발로 나누어진다. 지금까지 고축적식물에 의한 중금속 축적 기작은 다섯 가지의 특이 기작으로 알려져 있는데, 1) 뿌리세포의 중금속 흡수 증진, 2) 식물체 조직내의 중금속 이동성 향상, 3) 중금속의 무독화(detoxification) 및 격리(sequestration), 4) 토양-뿌리 경계면에서의 중금속 유효도 증진, 그리고 5) 중금속 오염토양으로의 능동적인 뿌리의 성장 등이 이에 속한다. 일반적으로 토양 중 낮은 중금속 유효도는 식물복원 기술의 현장 적용에 있어 제한요소로 간주된다. 이를 극복하기 위해서는 위에 기술된 다섯 가지 기작 중 고축적식물의 뿌리가 근권 토양중 중금속의 화학변화에 미치는 영향을 이해하는 것이 매우 중요하다. 식물 뿌리에 의한 근권 토양의 pH 변화와 뿌리에서 나오는 분자량이 적은 유기산(low-molecular-weight organic acids, LMWOAs)과 같은 유기성 분비물은 근권부 토양의 화학적 특성을 변화시키고 결과적으로 중금속의 유효도를 변화시킨다. 예를 들어 뿌리에서 나오는 $H^+$ 이온은 토양 pH를 감소시키고 이에 따라 중금속의 유효도는 증가한다. 또한 고농도의 중금속에 노출된 뿌리는 많은 양의 유기물질을 분비하게 되고 근권 토양에 축적되는 이 유기물질은 토양중 중금속과 결합하여 유기복합물질(organo-metallic complexes)을 형성하면서 유효도를 증가시킨다.
Much research has been conducted in the field of phytoremediation since the discovery of the range of plants known as hyperaccumulators. Research has focused simultaneously on elucidating the mechanism of metal(loid) accumulation and development of practical techniques to enhance accumulation effici...
Much research has been conducted in the field of phytoremediation since the discovery of the range of plants known as hyperaccumulators. Research has focused simultaneously on elucidating the mechanism of metal(loid) accumulation and development of practical techniques to enhance accumulation efficiency. To date, it is generally understood that there are five specific mechanisms employed by hyperaccumulating plant species that are either not or under utilized by non-hyperaccumulators. These include 1) enhanced metal(loid)s uptake through the root cell, 2) enhanced translocation in plant tissue, 3) detoxification and sequestration, 4) enhanced metal availability in soil:root interface, and 5) active root foraging toward metal(loid) enriched soils. Among these mechanisms, understanding of the plant-root effect on metal(loid) dynamics and subsequent plant uptake is vital to overcome the inherit limitation of phytoremediation caused by low metal(loid) solubility in soils. Plant roots can influence the soil chemistry in the rhizosphere through changes in pH and exudation of organic compounds such as low-molecular-weight organic acids (LMWOAs) which consequently change metal(loid) solubility. The decrease in soil pH by plant release of $H^+$ results in increased metal solubility. Elevated levels of organic compounds in response to high metal soil concentrations by plant exudation may also increases metal concentration in soil solution through formation of organometallic complexes.
Much research has been conducted in the field of phytoremediation since the discovery of the range of plants known as hyperaccumulators. Research has focused simultaneously on elucidating the mechanism of metal(loid) accumulation and development of practical techniques to enhance accumulation efficiency. To date, it is generally understood that there are five specific mechanisms employed by hyperaccumulating plant species that are either not or under utilized by non-hyperaccumulators. These include 1) enhanced metal(loid)s uptake through the root cell, 2) enhanced translocation in plant tissue, 3) detoxification and sequestration, 4) enhanced metal availability in soil:root interface, and 5) active root foraging toward metal(loid) enriched soils. Among these mechanisms, understanding of the plant-root effect on metal(loid) dynamics and subsequent plant uptake is vital to overcome the inherit limitation of phytoremediation caused by low metal(loid) solubility in soils. Plant roots can influence the soil chemistry in the rhizosphere through changes in pH and exudation of organic compounds such as low-molecular-weight organic acids (LMWOAs) which consequently change metal(loid) solubility. The decrease in soil pH by plant release of $H^+$ results in increased metal solubility. Elevated levels of organic compounds in response to high metal soil concentrations by plant exudation may also increases metal concentration in soil solution through formation of organometallic complexes.
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성능/효과
(1996) found that exposing several Ni hyperaccumulator species of Alyssum to Ni resulted in a large and proportional increase in the levels of histidine in the xylem sap, which was shown to be complexed with Ni. Furthermore, it was shown that supplying exogenous histidine to the nonaccumulator A. montanum, either as a foliar spray or in the root medium, considerably increased the nickel tolerance of this species and greatly increased nickel flux through the xylem.
Blaylock and Huang (2000a) also suggest that hyperaccumulators are able to modify the rhizosphere to enhance metal solubility. Their study showed increases in Zn accumulationin the shoot of the non-hyperaccumulator Thlaspi arvense, when it was co-cultivated with a Zn hyperaccumulator Thlaspi caerulescens.
(2001) investigated the difference in adsorption ability between a Cd hyperaccumulating ecotype, Thlaspi caerulescens, from Ganges in southern France and a Cd non-hyperaccumulating ecotype, Thlaspi caerulescence, from Prayon in Belgium. Their study showed that the Vmax for Cd influx was five fold higher in the high Cd ecotype than in the low Cd ecotype. This result also suggests a high-affinity Cd transport system in the Cd hyperaccumulating population of Thlaspi caerulescens which enabled it to accumulate a higher level of Cd than the non-hyperaccumulating ecotype.
(1996), this ability is associated with the amount of metal transporters in the root. Their study showed that the Zn hyperaccumulator Thlaspi caerulescens had a significantly higher uptake rate of Zn when compared to the non-hyperaccumulator Thlaspi arvense. The hyperaccumulation was attributed to the plasma membranes of root cells of Thlaspi caerulescens having a higher density of Zn transporters.
However, decreases in the soil solution pool accounted for only about 1%and 50% of the total Zn and Cd uptake, respectively, by the plants. These results suggested that most of the Zn and approximately 50% of the Cd taken up by Thlaspi caerulescens were from a non-labile fraction. This meant that either Thlaspi caerulescens was highly efficient at mobilising Zn orCd which was not initially soluble, or that the soils used had low buffering capacities that were able to replenish the soil solution Zn and Cd within a very short time.
In the root, Zn was mainly coordinated with histidine, while in the xylem sap Zn was mainly transported as a free hydrated cation, and in the leaves it was primarily complexed with organic acids. This study indicated that the role of metalinducing ligands seemed to depend on both metal type and plant species. Therefore, further work is required to ascertain whether metal-induced production of histidine (or perhaps other nitrogen-containing ligands) is found in other groups of nickel hyperaccumulators, or indeed in other hyperaccumulators for other metals.
후속연구
Even though several researchers have suggested the possible mobilizing ability of the hyperaccumulator root, a complete understanding of the exudation of phytosiderophores and organic acids, and the resulting changes in pH and redox potential, which are considered key factors in controlling metal availability in the rhizosphere (Lombi et al., 2000), isstill poorly understood and further studies will be of great benefit to the field.
Even though these studies show the possibility that metal-transporters are in operation in the plant roots, more studies with other hyperaccumulators and metals are necessary for confirmation of metal transporter function.
This study indicated that the role of metalinducing ligands seemed to depend on both metal type and plant species. Therefore, further work is required to ascertain whether metal-induced production of histidine (or perhaps other nitrogen-containing ligands) is found in other groups of nickel hyperaccumulators, or indeed in other hyperaccumulators for other metals.
, 2003). These contrasting findings may be due to the different soil and plant types employed in each study and hence further investigation considering the affect of plant and soil type on soil acidification would be of great benefit.
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