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Ecophyslc:)Logical EFFECTS of HIGH CONCENTRATIONS OF Eco p Hy slc:)LOGIcAL EFFECTS OF HIGH CONCENTRATIONS OF IRON AND OTHER HEAVY METALS ON ER I OPHO RUM ANGUSTIFOLIUM HON CR. AND PHRAGMITES AUSTRALIS (CAV ) TRIN EX STE.UDEL SUZANNE MARY MANSFIELD Deprtnient of Animal and Plant Sciences, University of Sheffield Thesis for the degree of Doctor of Philosophy JANUARY 1989 To rriu fari,i lt..i nd th rnflioru of G W.M E•:cOPHYSIOLOG]:cAL. EFFECTS F HIGH CONCENTRATIONS OF IRc)N AND OTHER HEAVY METAL.S o ERIOPHORUM ANGUSTIFOLIUM HONCR. AND lI:ri (CAV. ) TRIN. EX STEUDEL. S N Mansfield I)epartrrient of Animal and Plant Sciences, Universit y of Sheffield SUMMARY Little is known of the effect of hi g h conc:entrations of heavy metals on the growth of wetland plants, although it is has long been sUg cested that adaptation to waterlogged anaerobic environments involves the capacity to exclude dissolved iron. This work investigates the chemical dynarruics of selected metal toxins, in particular Fe in relation to concentration in the substrate and uptake b y Eriophorum anustifoliuff, and Phraqmites australis taken from mine populations (Parys Mountain, Anglesey and Crym].yn Bo g , Swansea) and a non-mine population (Skipwith Common, Yorks) and to provide inforrriation on the resistance of these two species to high concentrations of Fe and other heav y metals. Both field and laboratory work have been used to focus on the above areas. The first part of the study is based on the analysis of plants and soil samples collected from the field and relates heav y metal concentration in plant tissue to concentrations in soil/sediments. The second part of the study is based on laboratory work to determine individual factors influencing heav y metal uptake and resistance in the study species. Fe, Mn, Cu, Zn and Pb concentrations in plant tissues reflected those in the soil. Seasonal fluctuations in metal concentration in plant tissues were observed but the bulk differences were related to soil heterogeneity between sites. In culture solutions, Fe uptake in both species increased with increasin g Fe supply. E. anqstifalium frorri the Parys site was less sensitive to hi g h Fe concentrations relative to plants from the Skipwith site. Fe-uptake by E angustjfol ium, was stron g l y influenced by pH. Fe--uptake by plants was uneffected by the presence of Fe-plaques on roots. A relationship between Fe and P is highlighted enhancing P availability to plants of E. angjtifolium in th presence of 100 mg /l Fe by treatin g half the roots with Fe and the remaining half with 0.1-strength Rorison solution with 1 mg/l Fe, stimulated root oxidation of Fe and the accumulation of Fe-plaque, reducing translocat ion of Fe to shoots. P].ants of the same species with Fe-plaque an the roots accumulated more P than plants witI-,out I::ilaque, the bulk of thi.s P was immobile. E. an q ustifolium from the Parys Mountain site was found to be rriore tolerant to Cu but more sensitive to Mn, the reverse was true for p).ants from the Skipwi.th site. I The irnp1itions of this work in reition to plant qrowth and resistarie to heavy metal—enriched environments are discussed. It is suqqested that both species rriaq be constitutionallj tolerant to Fe and Mn. Plants from the Parus site mat.j he more resistant to high concentrations of Cu, Zn and Pb. A CR N C) WI ..Er) GEM E NT S I am grateful to Dr. B. D. Wheeler and Dr. A.. J. M.. Baker for their advice and support throughout this research.. Many thanks are also due to Dr. B. C.. Jarvis for providing supervision and advise for Chapter 6. I would like to thank Professor D. H. Lewis for providing research facilities and the Natural Environment Research Council for financial support.. Thanks are also due to the technical staff, Miss R. Cook, Mr. J. Re]].u, Mr.. A. Fairburn, Mr. B. Keen, Mr. R. P.radlej, Mr G. Woods and the Workshop for their everwilling advice and technical assistance and to Mr.. T. Croft. Mu thanks to all in C41 for their friendship and invaluable discussions, to those WhO have not been mentioned but who have made this Ph.D possible and to those who have enriched rrILj time at the department of Animal arid Plant Sciences. Finally, but not leastly, I am forever indebted to John without whose support and constant encouraernent would have made the last few months much harder ECOPHYSIOLOGICAL EFFECTS OF HIGH CONCENTRATIONS OF (RON AND OTHER HEAVY METALS (::)N ER I OPHORUM ANGUST I FOL i UM HONCK, ANt) PI-IRAGMITES AUSTRALIS(CAV. ) TRII\l EX STEUDAL CONTENTS Paqe SUMMARY L A CKNOWLEDGEMENTS /i CHAPTER 1 GENERAL INTRODUCTION 1.1 Sail contamination bj 1. heavLi rrietals and the evolution of tolerance in plants 1.2 Response of 4 terrestrial plants to toxic metals 1.3 Wetland plants and 9 toxic metals :1.4 Aims and Approaches Chapter 2 SOIL SEDIMENT HEAVY METAL CONCENTRATIONS AND CORRESPONDING METAL CON CENT RAT I (INS IN PLANT TISSUES FROM A CONTAMINATED AND UNCONTAMINATED WETLAND: A PILOT STUDY 2.1 Introduction 25 2.1.1 Ecoloqi of Eriophorum 36 an q ustifolium and Phraqmites austral is 2.1.2 Site discriptions 46 22 Material and Methods 50 2.3 Results 65 2.4 Discussion 72 2.5 Conc:lusions 92 CHAPTER 3 SEASONAL DYNAMICS IN HEAVY METAL CONCENTRAT I (INS IN CONTAM I NATED AND UNCONTAMINATED WETLANDS 3.1 Introduction 96 3.2 Materials and methods 12 3.3 Results 105 34 Discussion 115 3.5 Conclusions 127 CI-IAPTER 4 THE EFFECT OF INCREASING SUPPLY OF SELECTED HEAVY METALS ON UPTAKE CONCENTRATION IN TISSUES AND GROWTH OF ERIOPHORUM ANGUSTIFOLIUM ANI) PI-IRAGM lIES AUSTRALIS 4.1 Introduction 130 4.2 Material and methods 137 4.3 Results 145 4.4 Discussion 159 4.5 Conclusion 182 CHAPTER 5 EFFECT OF NUTRIENTS ON IRON UPTAKE AND GROWTH OF ER I OPHORUMANGUST I FOL I UN AT DIFFERENT CONCENTRATIONS OF IRON 5.1 Introduction 185 5.2 Materials and methods 190 5.3 Results 198 5.4 Discussion :24 5.5 Conclusions 219 CHAPTER 6 FACTORS INFLUENCING THE UPTAKE OF IRON BY ERIOPHORUM ANGUST I FOL I UN UNDER CONTROLLED CONI) IT I ONS 6.1 Introduction 221 6.2 Materials and methods 226 6.3 Results 235 6.4 Discussion 241 6.5 Conclusions 251 CHAPTER 7 GENERAL DISCUSSION 7. 1 Mechanisms of metal 256 resistance in Eriophorum anustifolium and Phracimites australis 7.2 The mechanisms of 267 tolerance of ErioEhorum anciustifolium and Phra q mites australis to toxic concentrations of iron and other metals 7.3 EcotLipic differentiation 278 i n Eriophorum anciust i foi 1LUTI and Phraqniites austral is 7.4 :[nterpretation of the 290 work in relation to the types of tolerance shown n arciustifoljurn and Phrarnites_australis 7.5 Interpretation of the work 293 in relation to plant resistance to anoxic con dit ions 131 EL IOGRAPHY 296 APPEND I CES I. Composition of 315 Rorisons solution II Propaqation of 316 plants for solution culture work III. Nitric-perchloric 319 diaestion method and colorimetric method for determininq phosphus IV Seasona]. 234 fluctuations in element composit ion in plant tissues additional data (Chapter 3) V. Construction of a 332 quench curve for countinq 59Fe VI. Results of 334 reciprocal transplantation of Phraqmi tes australis from Parys Mountain and Skipwith Common, April 1986 Chapter 1 General Introduction 1.1 Soil contamination bq heavq metals and the evolution of tolerance in plants The heavq metals, defined bq Passow, Rothstein & Clarkson (1961), are those rrietals that have a densit!J greater then 5 and, as a group, contain about 38 elements, including iron (densitu 79 q/cm3, Gilmore 1979), Mn, Cu. Zn and Pb. Their cornrrton features in relation to plant growth is that in excessive quantities they are toxic causing, for example root stunting and ultimatel y death (Bradshaw, McNeilly & Gregory 1965). Local enrichment of heavy metals in soils, either through natural processes (geochemical anomalies) or human activities (exploitation of mineral resources), is usually 1 associ.ated with increased resistance levels in plants (Antonovics, Bradshaw & Turner 1971). Thus, the presence of a species or a race on metal-contaminated soi].s sugciests that it is tolerant to metal toxicitt. Such edaphic adaptation, however, necessitates a specialized phqsioloqq resultinq in either constitutional tolerance within the species or ecotpic differentiation of the species, into metal--tolerant races which are specificallu adapted phqsiologicallt (Baker 1981). According to P.radshaw (1952), the evolution of metal tolerance in plant populations arises as a result of selection in localized areas of heavq metal contamination. Metal-tolerant populations contain individuals which have been selected and which are able to tolerate much hiqher concentrations of metals than individuals qrowinq in non-contaminated areas (Antonovics etal. 1971). The mechanism of tolerance can be specific such that possession of tolerance to one or two metals does not neccessarilq confer tolerance to another metal not present at the site in high concentrations (Ernst 1976, Qureshi, Thurman, Hardwick & 2 Collin 1995). 1.1.1 Strategies for survival in metal-contaminated soils Levitt (199w) has described the nature and meaning of stress injury and resistance in organisms. In accordance to Levitt's descriptions, heavy metals in the plant environment can operate as 'stress' factors in that they can cause 'strain' by affecting physiological processes in plants. In so doinci they can reduce viqour, or totally inhibit plant growth. 'Sensitivity' describes the effects of stress resulting in injury or death of the plant, while 'resistance' refers to the reaction of a plant to heavy metal stress in such a way that it can grow and reproduce successfully.
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