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Soil~Plant Relationships: An Ecological Approach PDF

297 Pages·1987·7.708 MB·English
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SOIL -PLANT RELA TIONSHIPS An Ecological Approach DAVID W. JEFFREY CROOM HELM London & Sydney TIMBER PRESS Portland, Oregon © 1987 David W. Jeffrey Croom Helm Ltd, Provident House, Burrell Row, Beckenham, Kent BR3 lAT Croom Helm Australia, 44-50 Waterloo Road, North Ryde, 2113, New South Wales British Library Cataloguing in Publication Data Jeffrey, David W. Soil-plant relationships: an ecological approach. 1. Botany - Ecology I. Title 581.5'26404 QK901 ISBN-13: 978-0-7099-1464-8 e-ISBN-13: 978-94-011-6076-6 DOl: 10.1007/978-94-011-6076-6 Pbk First published in the USA 1987 by Timber Press, 9999 S.W. Wilshire, Portland, OR 97225, USA All rights reserved ISBN -13: 978-0-7099-1464-8 Contents Preface vii Part I: A plant-centred biological complex 1 1 Plants, roots and ion absorption 3 2 Mineral composition of plant tissues and the function of ions 18 - 3 Plants and water 50 4 Symbiotic and other associations for nutrient capture 63 - 5 Herbivores, decomposers and other soil organisms 82 6 Vegetation and fire 91 Part ll: Environmental complexes 95 7 Soil formation 97 8 Soil matrix and soil water 109 9 Soil atmosphere and soil temperature 129- 10 Some examples of mineral nutrient supply 136- 11 Measuring availability of nutrients and toxic ions 150 12 Experimental approaches to the study of soil variables 161 Part ill: Interactions in the real world. Some case histories 173 13 The autecology of two contrasting species 175 14 Restoration of derelict land 185 15 Two aspects of forest mineral-nutrient economy 202 16 Australian heathlands and other nutrient-poor terrestrial ecosystems 211 17 Three aspects of the Alaskan Arctic tundra complex 224 18 Saltrnarshes and the coastal zone 235 v CONTENTS 19 Calcareous and serpentine soils and their vegetation 257 Further reading 277 Bibliography 280 Index 291 vi Preface Soil-plant relationships once had a limited meaning. To the student of agriculture it meant creating optimum conditions for plant growth. To the ecologist it meant explaining some plant community distribu tion patterns by correlation with soil type or conditions. This dual view has been greatly expanded at an academic level by the discovery of the ecosystem as a practical working unit. A flood of concepts and information subsequently emerged from the International Biological Programme. At a totally different level of resolution, it is appreciated that certain soil-based ecological problems have a molecular basis, and must be addressed by physiological or biochemical approaches. From ecosystem to molecule we have powerful new tools to increase the flow of ecological data and process it for interpretation. Society is now experiencing a series of adverse global phenomena which demand an appreciation of soil-plant relationships. These include desertification leading to famine, soil degradation accom panying forest destruction, acidification of watersheds and the spasmodic dispersal of radionuclides and other pollutants. It is public policy, not merely to identify problems, but to seek strategies for minimising their ill effects. This book is written as a guide to soil-plant relationships, cen trally oriented towards ecology, but of interest to students of geo graphy and agriculture. For ecology students it will bring together subfields such as microbiology, plant physiology, systematics and pro vide interfaces with animal biology, meteorology and soil science. Ideas contributing to the formulation of hypotheses are emphasised, rather than overloading readers with information better obtained first hand from the primary literature or encyclopaedic reviews. The selection of case histories in ecological investigation indicates the range of possible approaches to demonstrate that ecology is a positive and applicable science, capable of earning its keep in the eyes of the world. I must gratefully acknowledge the contributions made to my ideas by past and present colleagues in Ireland, Britain, Australia and North America. They are too numerous to name individually and their generosity has been unbounded. However, lowe most in terms of concept development and stimulation to my students and my family. David W Jeffrey Dublin 1986 vii Part One A plant-centred biological complex The theme of this section is the response of plants in relation to ions and water in the soil environment. It draws special attention to the close biological relationships between plants, microorganisms and other organisms in the food web. The role of plants as primary producers is taken for granted, even though limits to plant growth imposed by soil conditions will be encountered many times. This account should be read in conjunction with the prolific literature on production ecology. 1 1 Plants, roots and ion absorption INTRODUCTION The view of ecology adopted here is that it comprises the study of ecosystems and their surroundings. Plants are thus seen in relation to the cluster of organisms most closely associated with them and to their environment. Plants are located in an environmental complex of energy, atmosphere and soil. Within an ecosystem context, plants have three principal functions: (a) Absorption of photosynthetically active radiation which is applied to the synthesis of carbon-to-carbon bonds. Compounds synthesised in this way serve as the only energy currency for the ecosystem. (b) Absorption and assimilation of ions as a source of essential elements for the ecosystem. Photosynthesis is necessary for ion absorption and assimilation, and ions are an essential part of the photosynthetic system and all other metabolism. (c) Plants process water on a sufficiently large scale to make a large contribution to the hydrological cycle of the Earth. Some of this water use is ancillary to photosynthesis, general metabolism and the fabric of plant structure. Much is involved with the special place of plants in unavoidably intercepting solar heat while simultaneously optimising gas exchange. Leaves are cooled by evaporative water loss. These activities are represented in a simple way by Figure 1.1, which symbolises leaves and roots configured for their respective functions. Storage of energy and ions is also regarded as a universal feature. 3 PLANTS, ROOTS AND ION ABSORPTION Figure 1.1: Plants are organisms that are rooted in soil and receive and process energy, water and ions. They are the foundation of ecosystems and contribute to hydrologic and geochemical processes ENERGY Atmosphere Soil WATER The ecologist should observe the vegetation being studied with the eye of the designer, the systems analyst, the civil engineer and even the artist. This appraisal will prove valuable in hypothesis building. The simple realisation formalised by Raunkier, that plants' life forms can be readily classified and that whole communities have common morphologies and growth behaviours, is still important. It makes us realise for a start that sheer maximisation of biomass pro duction is not a characteristic of predominant ecological importance. Long-term persistence of a species is much more subtle, entailing the optimisation of: assimilating the materials for primary produc tion; producing and disseminating propagules; accommodating to the short-term variations in environment; accepting predation; and adapting to long-term environmental fluctuation and even cataclysm. The means for long-term persistence under particular soil environment regimes entails a range of features including 4 PLANTS, ROOTS AND ION ABSORPTION morphological and physiological plasticity, ecotype formation, species characteristics and indeed the collective properties of larger taxa. Ecologists will broadly recognise the association between the vetches and peas (Papilionoideae) and nitrogen fixation; between the heathers (Ericaceae) and survival on infertile, acid substrates; between the glassworts and seablites (Chenopodiaceae) and suc culence and salt tolerance; and between the Cactaceae and survival in arid zones. It seems overwhelmingly obvious that the vascular plants have, at least in part, had their evolution moulded by soil characteristics. ROOTS The three functions of all roots which are outwardly directed'to the environment, and which are worth discussing in an ecok>gical context, are absorption, anchorage and storage. Storage may, according to some opinions, fall into the category of internal metabolic functions for roots, which include, for example, the pro duction of growth regulators such as kinetin and abscisi~ acid (Russell 1977). However, it is not possible to discuss the absqrption and fate of mineral metabolites without reference to storage~ Root growth and root structure Unlike the complex apex of the shoot, the root apex is small and compact. It cuts off files of cells which differentiate to produce epidermis, cortex and vascular bundles behind the apex and cells of the root cap before it. The first stage of cell differentiation is elonga tion. The energy for this process is generated from the translocation of osmotically active metabolites, sugars or ions, to the vacuoles of the expanding cells. Entry of water and development of a substantial pressure potential elongates the cells. Orientation of cellulose fibrils in cell walls, which resembles that of the hoops of a barrel, co ordinates the direction of growth. This uniaxial extension growth can be thought of as a jacking action, with a force of up to 10 bars (1.0 MPa) , which drives the root cap through the soil, reacting against the frictional and other forces gripping the older parts of the root. The dimensions of a typical root apex, in the order of 0.2 mm diameter, are sufficiently small to penetrate most inter-aggregate 5

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