. Uptake of phytotoxic amounts of metal by higher plants or algae can result in inhibition of several enzymes, and in increase in activity (= induction) of others. Two mechanisms of enzyme inhibition predominate: (1) binding of the metal to sulphydryl groups, involved in the catalytic actionor structural integrity of enzymes, and (2) deficiency of an essential metal in metalloproteins or metal-protein complexes, eventually combined with substitution of the toxic metal for the deficient element. Metal accumulation in the cellular compartment of the enzyme is a prerequisite for enzyme inhibition in vivo. The induction of some enzymes is considered to play a significant role in the stress metabolism, induced by metal phytotoxicity. Peroxidase induction is likely to be related to oxidative reactions at the biomembrane; several enzymes of the intermediary metabolism might be stimulated to compensate for metal-sensitive photosynthetic reactions. The induction of enzymes and metal-specific changes in isoperoxidase pattern can be used as diagnostic criteria to evaluate the phytotoxicity of soils, contaminated by several metals. Lines for future research on metal phytotoxicity are proposed, involving the study of inhibition and induction of enzymes at the different cell membranes (especially the plasmamembrane) in vivo.

Effects of metals on enzyme activity in plants

http://directory.umm.ac.id/Data%20Elmu/jurnal/P/PlantScience/PlantScience_Elsevier/Vol157.Issue1.2000/5399.pdf

Antioxidative parameters in the seedlings of pigeonpea (Cajanus cajan (L.) Millspaugh) in response to Zn and Ni stresses.

Heavy metal pollution is one of the most troublesome environmental problems faced by mankind nowadays. Copper, in particular, poses serious problems due to its widespread industrial and agricultural use. Unlike other heavy metals, such as cadmium, lead, and mercury, copper is not readily bioaccumulated and thus its toxicity to man and other mammals is relatively low. On the contrary, plants in general are very sensitive to Cu toxicity, displaying metabolic disturbances and growth inhibition at Cu contents in the tissues only slightly higher than the normal levels. The reduced mobility of Cu in soil and sediments, due to its strong binding to organic and inorganic colloids, constitutes, in a way, a barrier to Cu toxicity in land plants. In aqueous media, however, plants are directly exposed to the harmful effects of Cu and, thus, algae and some species of aquatic higher plants are more easily subjected to Cu toxicity. Excess Cu inhibits a large number of enzymes and interferes with several aspects of plant biochemistry, including photosynthesis, pigment synthesis, and membrane integrity. Perhaps its most important effect is associated with the blocking of photosynthetic electron transport, leading to the production of radicals which start peroxidative chain reactions involving membrane lipids. Copper effects on plant physiology are wide ranging, including interference with fatty acid and protein metabolism and inhibition of respiration and nitrogen fixation processes. At the whole plant level Cu is an effective inhibitor of vegetative growth and induces general symptoms of senescence. The high toxicity of Cu to plants has led to the evolution of several strategies of defense. Among the most important ones is the production of Cu-complexing compounds. Although the nature, structure, and function of these compounds is still controversial, they can be divided into two main groups: metallothionein-like compounds and phytochelatins. The latter appears to constitute the most widespread response of plants to stresses provoked by metals, including Cu.

Biochemical, physiological, and structural effects of excess copper in plants

Nearly 60 years ago, Prat (1934) initiated the research of heavy metal resistance in plants when he was analysing the growth performance of two populations of Silene dioica (Melandrium sylvestre), one from a copper mine and one from a non-mine soil. He was able to demonstrate a heritable copper resistance in the mine population, relative to the non-mine population, which he explained as a result of evolution by natural selection. Nearly 20 years later Bradshaw (1952) and Baumeister (1954) started further research on ecological and physiological differentiation between plants from metal-enriched and noncontaminated habitats. The species chosen for study were predominantly Agrostis capillaris in the Bradshaw group (Jowett 1959; Gregory 1965; McNeilly 1965; Antonovics 1966) and Silene vulgaris in the Baumeister group (Broker 1962; Ernst 1964; Gries 1965; Riither 1966). In the late 1950s Duvigneaud (1958), while studying the vegetation on metalliferous soils in Central Africa, added to the above approaches a phytogeographic one and introduced the study of speciation processes in metallophytes. In the 1950s, the study of evolutionary and physiological aspects of metal resistance was hampered by the absence of convenient techniques for measuring metal concentrations in small plant samples. The techniques available for metal analysis were either timeconsuming, such as phase separation (Ernst 1964), or costly and only applicable for laboratory-raised plant material, i.e. radiolabelling (Turner & Gregory 1967; Peterson 1969). Only after applying atomic absorption spectrophotometry on wet-ashed plant material (Reilly 1967) did time and cost-effective metal analyses become possible.

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Metal tolerance in plants

Comparative studies of H2O2 detoxifying enzymes in green and greening barley seedlings under cadmium stress

This paper provides information on the effects of toxic concentrations of cupric sulphate on the growth of lettuce (Lectuca sativa) seedlings. Root growth is completely inhibited at 5 x 10-2M and germination stops altogether at 10-1M. The relative inhibition of root growth is stronger than that of hypocotyl growth. Various metabolites and hormones are partially capable of relieving copper inhibition.



Catalase, peroxidase and IAA-oxidase activity shows increments directly proportional to the concentration of copper. It is obvious that growth is inversely proportional to enzyme activity. The increased level of these enzymes is probably due to an accelerated protein synthesis.

Characterization of Copper Texicity in Lettuce Seedlings

Growth and metabolism of germinating rice (Oryza sativa L.) seeds as influenced by toxic concentrations of lead

The present experiment provides information on the phosphorus compounds in rice seeds and elucidates the changes they undergo during germination. In ungerminated seeds, the bulk of total-P appears in phytin (about 76 per cent). It is then dephosphorylated in course of germination with a simultaneous accumulation of large amounts of inorganic-P. Lipid-P increases very rapidly from 0 to 24 hours. The increase up to 72 hours is gradual, after which it drops at 96 hours and again rises to a maximum after 120 hours. The ester-P and RNA-P, fractions increase in concentration with time of germination (except 120 hours). Protein-P begins to fall after 48 hours, while DNA-P remains more or less constant throughout the experiment.



The two pH optima recorded for phytase activity at 4.0 and 9.0, suggests that there exist two phytases, one acidic and the other alkaline. Both behave similarly during germination with a continuous increase throughout the course of the experiment. The enzyme with an optimal pH at 4 hydrolyses phytin more actively than the other with the pH optimum at 9.0. Phytase shows maximum activity at a stage when most of the phytin has disappeared; the metabolic significance is uncertain.

Changes in Phosphorus Fractions and Phytase Activity of Rice Seeds during Germination

Effects of Toxic Concentrations of Copper on Growth and Metabolism of Rice Seedlings

Assay of the hydrolyzing enzyme levels, viz.α-amylase, adenosine triphosphatase (ATPase), phytase and ribonuclease (RNase) in the germinating Mungbean (Phaseolus aureus) seeds is made to evaluate the degree of substrate breakdown at different periods of germination. In addition to the cotyledons, the principal site of hydrolases, an evidence of their occurrence in the growing embryo has been presented. Enzyme activities in storage tissues, i. e. cotyledons and growing embryo, i. e. axis rise to a maximum sometime between initial and final phase of germination and then declined.

S. Mukherji

Papers

Characterization of Copper Texicity in Lettuce Seedlings

Growth and metabolism of germinating rice (Oryza sativa L.) seeds as influenced by toxic concentrations of lead

Changes in Phosphorus Fractions and Phytase Activity of Rice Seeds during Germination

Effects of Toxic Concentrations of Copper on Growth and Metabolism of Rice Seedlings

Enzyme activities in germinating Mungbean (Phaseolus aureus) seeds and their relation with promoter and inhibitors of protein synthesis