Plant physiology (biological and medical physics, biomedical engineering)

Biological and Medical Physics, Biomedical Engineering Maria Duca Plant Physiology Tai Lieu Chat Luong Plant Physiology BIOLOGICAL AND MEDICAL PHYSICS, BIOMEDICAL ENGINEERING The fields of biological and medical physics and biomedical engineering are broad, multidisciplinary and dynamic They lie at the crossroads of frontier research in physics, biology, chemistry, and medicine The Biological and Medical Physics, Biomedical Engineering Series is intended to be comprehensive, covering a broad range of topics important to the study of the physical, chemical and biological sciences Its goal is to provide scientists and engineers with textbooks, monographs, and reference works to address the growing need for information Books in the series emphasize established and emergent areas of science including molecular, membrane, and mathematical biophysics; photosynthetic energy harvesting and conversion; information processing; physical principles of genetics; sensory communications; automata networks, neural networks, and cellular automata Equally important will be coverage of applied aspects of biological and medical physics and biomedical engineering such as molecular electronic components and devices, biosensors, medicine, imaging, physical principles of renewable energy production, advanced prostheses, and environmental control and engineering Editor-in-Chief: Elias Greenbaum, Oak Ridge National Laboratory, Oak Ridge, Tennessee, USA Editorial Board: Masuo Aizawa, Department of Bioengineering, Tokyo Institute of Technology, Yokohama, Japan Judith Herzfeld, Department of Chemistry, Brandeis University, Waltham, Massachusetts, USA Olaf S Andersen, Department of Physiology, Biophysics and Molecular Medicine, Cornell University, New York, USA Mark S Humayun, Doheny Eye Institute, Los Angeles, California, USA Robert H Austin, Department of Physics, Princeton University, Princeton, New Jersey, USA James Barber, Department of Biochemistry, Imperial College of Science, Technology and Medicine, London, England Howard C Berg, Department of Molecular and Cellular Biology, Harvard University, Cambridge, Massachusetts, USA Victor Bloomfield, Department of Biochemistry, University of Minnesota, St Paul, Minnesota, USA Robert Callender, Department of Biochemistry, Albert Einstein College of Medicine, Bronx, New York, USA Britton Chance, University of Pennsylvania Department of Biochemistry/Biophysics Philadelphia, USA Steven Chu, Lawrence Berkeley National Laboratory, Berkeley, California, USA Louis J DeFelice, Department of Pharmacology, Vanderbilt University, Nashville, Tennessee, USA Johann Deisenhofer, Howard Hughes Medical Institute, The University of Texas, Dallas, Texas, USA George Feher, Department of Physics, University of California, San Diego, La Jolla, California, USA Hans Frauenfelder, Los Alamos National Laboratory, Los Alamos, New Mexico, USA Ivar Giaever, Rensselaer Polytechnic Institute, Troy, NewYork, USA Sol M Gruner, Cornell University, Ithaca, New York, USA Pierre Joliot, Institute de Biologie Physico-Chimique, Fondation Edmond de Rothschild, Paris, France Lajos Keszthelyi, Institute of Biophysics, Hungarian Academy of Sciences, Szeged, Hungary Robert S Knox, Department of Physics and Astronomy, University of Rochester, Rochester, New York, USA Aaron Lewis, Department of Applied Physics, Hebrew University, Jerusalem, Israel Stuart M Lindsay, Department of Physics and Astronomy, Arizona State University, Tempe, Arizona, USA David Mauzerall, Rockefeller University, New York, New York, USA Eugenie V Mielczarek, Department of Physics and Astronomy, George Mason University, Fairfax, Virginia, USA Markolf Niemz, Medical Faculty Mannheim, University of Heidelberg, Mannheim, Germany V Adrian Parsegian, Physical Science Laboratory, National Institutes of Health, Bethesda, Maryland, USA Linda S Powers, University of Arizona, Tucson, Arizona, USA Earl W Prohofsky, Department of Physics, Purdue University, West Lafayette, Indiana, USA Andrew Rubin, Department of Biophysics, Moscow State University, Moscow, Russia Michael Seibert, National Renewable Energy Laboratory, Golden, Colorado, USA David Thomas, Department of Biochemistry, University of Minnesota Medical School, Minneapolis, Minnesota, USA More information about this series at http://www.springer.com/series/3740 Maria Duca Plant Physiology 123 Maria Duca University of Academy of Sciences of Moldova Chişinău Moldova ISSN 1618-7210 ISSN 2197-5647 (electronic) Biological and Medical Physics, Biomedical Engineering ISBN 978-3-319-17908-7 ISBN 978-3-319-17909-4 (eBook) DOI 10.1007/978-3-319-17909-4 Library of Congress Control Number: 2015939679 Springer Cham Heidelberg New York Dordrecht London © Springer International Publishing Switzerland 2015 This work is subject to copyright All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed The use of general descriptive names, registered names, trademarks, service marks, etc in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made Printed on acid-free paper Springer International Publishing AG Switzerland is part of Springer Science+Business Media (www.springer.com) Preface The past decades came with tremendous advances in understanding molecular systems that lie at the core of life itself, a fact which has revolutionized biological research and the field of plant physiology was not an exception Moreover, with the current advent of high throughput technologies in genomics and proteomics the potential appears to reveal the most subtle details regarding the molecular actors and the processes in which they are involved But for being able to interpret and make use of such complex data, to understand its place and significance in the global context of plant metabolism, one must first hold basic knowledge of the key processes in the life of the plants, integrated across several dimensions like structure, function, ecology, etc Plant physiology can offer such an integrated view The subject of plant physiology is highly interdisciplinary and builds upon the knowledge derived from fields like botany, zoology, plant morphology and anatomy, cytology, biochemistry, molecular biology, etc While at the theoretical level one of the priorities is to integrate the information from these scientific areas for a most complete understanding of the processes undergoing in living system, at the practical level this field comes with abundant experimental knowledge and wellestablished practices inherited from previous decades that allow to manipulate crop species in the desired manner, even if the theoretical aspects are not always completely elucidated The course, presented by this book, offers the possibility to enter into the essence of the most important phenomena of the living matter—photosynthesis, respiration, growth and development, etc By being conceived in agreement with the requirements of modern biology, Plant Physiology offers a perspective over the instruments and methods which allow the manipulation of the vegetal organism and which lie at the foundation of biotechnology as we know it today The present book is not one that reflects only the principles and fundamental directions of plant physiology by using the scientific literature passed through the prism of own reflections, but also includes results of the personal research summarizing a big volume of experimental data v vi Preface The presented content adheres to the principle of applicability of the provided knowledge which means that theoretical topics are accompanied by real examples of their relevance from agriculture, plant breeding, etc A special place is left for graphical illustrations, diagrams, pictures, which occupy a significant proportion of the content and are meant to facilitate the process of assimilating the information The author wants to thank the university professor, habilitated doctor A.I Derendovschi for the detailed analysis of the content of the book and for the useful and constructive suggestions I am grateful and want to thank everyone who made a contribution to the appearance of this book—PhDs in Biology Angela Port, Ana Căpăţână, Aliona Glijin, Ana Bârsan, Elena Savca, Alexei Levitchi, Victor Lupascu, Ph.D students Lucia Ciobanu and all other students who helped me conceive this book I would like to thank Prof V Ciobanu, Prof V Reva, PhDs Elena Muraru, Tatiana Homenco, Otilia Dandara for the important suggestions regarding the undertaken approach and the full and complex support offered in the process of preparing and editing this book For the help provided in obtaining and consulting the most up to date scientific literature, I would like to thank my colleagues from the University of California, Riverside (USA)—Professors Isgouhi Kaloshen, Carol Lovatt, Seymour Van Gundy I would also like to express special gratitude to my family for the patience and understanding that they showed all these years Chişinău Maria Duca Contents 12 13 15 18 21 23 27 27 31 37 39 41 43 45 46 49 49 50 50 Introduction to the Educational Course of Plant Physiology 1.1 The Definition and Scope of Plant Physiology 1.2 Purposes of Plant Physiology as a Science 1.3 Research Methods Used by Plant Physiology References Plant Cell Physiology 2.1 The Cell as a Structural, Morphological, Functional Unit of Living Organisms 2.2 Structural Organization, Chemical Composition and Function of the Cell Wall 2.3 Structure and Ultrastructure of Cell Protoplasm 2.4 Structure and Function of Biological Membranes 2.5 Exchange of Substances Between the Cell and the Medium 2.5.1 Ion Flow into the Cell 2.5.2 Water Flow into the Cell References Water Regime 3.1 Role of Water in Plants 3.2 Water Content and State in Plants 3.3 Forms of Water in the Soil Accessible and Inaccessible Water 3.4 The Root System as a Specialized Organ for Water Absorption 3.5 The Influence of External Factors on Water Absorption Intensity 3.6 Water Elimination Physiological Importance of Plant Transpiration 3.6.1 Indices of Transpiration 3.7 Structure of the Leaf as an Organ of Transpiration vii viii Contents 3.8 Stomatal and Cuticular Transpiration 3.8.1 Stomatal Transpiration 3.8.2 Cuticular Transpiration 3.9 Water Absorption Mechanism and Ways of Its Circulation in Plants 3.9.1 Water Transport in Plants 3.10 Ecology of the Water Regime in Plants References 51 51 54 54 55 58 63 65 68 70 Photosynthesis 4.1 Importance of Photosynthesis and the Global Role of Green Plants 4.2 The Leaf as a Specialized Photosynthesis Organ 4.3 The Structure, Chemical Composition, Function and Origin of Chloroplasts 4.4 Photosynthesis Pigments 4.5 Photosynthesis Energetics 4.6 Photosynthesis Mechanism 4.6.1 Light Phase of Photosynthesis 4.6.2 The Dark Phase of Photosynthesis 4.7 Photorespiration 4.8 Endogenous Regulatory Elements of Photosynthesis 4.9 Ecology of Photosynthesis References 72 77 81 86 87 101 107 110 117 122 Respiration General Notions of Respiration Respiratory Enzymes A.N Bach’s and V.I Palladin’s Theories Respiration Mechanism 5.4.1 Genetic Link Between Respiration and Fermentation 5.4.2 Glycolysis—The Anaerobic Phase of Respiration 5.4.3 Krebs Cycle (Tricarboxylic Acid Cycle) 5.4.4 The Electron Transport Chain and the Energetic Outcome of Aerobic Respiration 5.5 Different Types of Respiratory Substrate Oxidation 5.6 Ecology of Respiration 5.7 Regulation and Self-regulation of the Respiration Process References 123 125 128 130 131 131 132 135 138 140 143 144 148 Mineral Nutrition of Plants 6.1 Importance of Mineral Elements in Plant Nutrition 6.2 Chemical Composition of the Ash 149 151 153 Plant 5.1 5.2 5.3 5.4 Contents ix 6.3 6.4 Methods of Mineral Nutrition Research The Root System as an Organ for Absorption and Transport of Mineral Elements 6.5 Physiological Role of Macroelements 6.5.1 Absorption, Transport and Metabolism of Nitrogen 6.5.2 Absorption, Transport and Metabolism of Sulfur 6.5.3 Absorption, Transport and Metabolism of Phosphorus 6.5.4 The Physiological Role of Other Macroelements 6.6 Physiological Role of Microelements 6.7 Mechanism of Absorption and Transport of Ions in Plants 6.7.1 Mineral Element Absorption 6.7.2 Mineral Element Transport 6.8 Soil as a Substrate for Plant Nutrition 6.9 Influence of Various Environmental Factors on Mineral Nutrition in Plants References Plant Growth and Development 7.1 The Concept of Plant Growth and Development 7.1.1 Dormancy in Plants (Repose) 7.2 Types of Plant Growth 7.3 Phases of Cell Growth and Development 7.4 Phases of Plant Growth and Development 7.5 Genetic Aspects of Plant Morphogenesis 7.6 Endogenous Factors of Plant Growth and Development 7.6.1 Auxins 7.6.2 Gibberellins 7.6.3 Cytokinins 7.6.4 Abscisic Acid 7.6.5 Ethylene 7.7 Photoperiodism and Yarovization 7.8 The Influence of Exogenous Factors on Plant Growth and Development 7.9 Plant Growth Movements—Tropism and Nasties 7.10 Self-Regulation of Plant Growth and Development References Plant 8.1 8.2 8.3 8.4 Biorhythms Classification and Mechanisms of Biological Rhythms Biological Rhythms in Plants Circadian Rhythms in Plants The Molecular Mechanism of the Circadian Clock 8.4.1 Environmental Signals Involved 154 155 156 156 165 168 170 175 178 178 181 182 183 184 187 189 190 193 193 195 197 199 203 207 210 213 215 218 221 222 225 228 231 233 236 239 241 242 10.8 Metabolism of Pollutants in Plants 301 Air pollution affects differently the various species of trees and lower plants Lichens, trees like the elm, the ash tree and the pine are so sensitive to SO2 that may serve as biological indicators of air pollution with this substance Walnut and apple trees are very sensitive to the harmful substances released into the atmosphere In these species drying of the tips of the branches and early flower detachment can happen In Quercus petraea and the black poplar Populus nigro significant defoliation and massive drying of the branches occur Pelargonium zonale because of its increased sensitivity to pollutants, can serve as an indicator of SO2 in the air by presenting a yellowing of leaves, defoliation, pronounced chlorosis of the leaf blade (at first on the edges, then progressing inwards along the leaf nerve) In Iris germanica, Vitis vinifera, Lotus corniculatus burns on the leaves and leaf necrosis caused by SO2 can be noticed which by reacting with water, produce H2SO3 and H2SO4 Plant species such as tobacco are sensitive to oxidants and leaves of cabbage (Brassica)—to aromatic polycyclic hydrocarbons, as well as lead, iron, chlorine Plants have an important role in cleaning the soil Due to the absorption capacity of the root system, a decrease in the content of all types of pollutants in the soil happens The plants are able to absorb from the soil nitroso compounds, oxides, pesticides, heavy metals, etc Uptake of pollutants by plants occurs also in the aquatic environment Aquatic plants are involved in water purification and absorption of biogenic elements, of hydrocarbons, phenols In the Netherlands Scirpus is widely used for water purification An active water purifier is Elchornia crassipes, which absorbs salts of heavy metals, insecticides and detergents Lemna absorbs heavy metals, nitrogen, phosphorus, limiting their spread 10.9 Biochemical Mechanism of Pollutant Transformation in Plants Transformation of pollutants in glycosides and elimination of xenobiotics is performed by compartmentalization of the metabolites in certain cellular structures (organelles) In the process of metabolizing xenobiotics degradation occurs by means of oxidation, reduction, decarboxylation, hydrolysis, etc reactions by means of conjugation or biosynthesis reactions By conjugation reactions some substances (amino acids, glucuronic acids) or some radicals (methyl, acetyl, sulfate, etc.) are introduced in the initial compound Enzymes that catalyze the metabolism of xenobiotics are localized in the cytoplasm, mitochondria and predominantly in the endoplasmic reticulum (microsomes) Thus, selectivity of herbicide action on different plants is owing to the specific enzyme system found in a particular species 2,4-dichlorophenoxyacetic acid, also known as the 2,4-D herbicide is absorbed by all sprayed plants However crops species, especially cereals have the ability to 302 10 Physiology of Plant Resistance to Unfavorable Environmental Factors Fig 10.15 The influence of 2,4-dichlorophenoxyacetic acid on cotton seedlings easily break down the side chain of the herbicide 2,4-D, which leads to its rapid metabolization, whereas weeds are perishing as a result of accumulating the 2,4-D herbicide, not due to its too high toxicity, but rather due to the excessive growth of plants in a short time Crop species can detoxify from the 2,4-D herbicide either by esterification of its carboxyl group with a monosaccharide either by forming a peptide bond with glutamic acid Detoxification is achieved also by hydroxylation of the aromatic nucleus, followed by a reaction of glycosidation, or by oxidation of the side chain If the side chain of the herbicide is removed, it loses its hormonal activity and 2,4dichlorophenol is formed as an intermediate product, which by means of natural glycosidation turns into the corresponding glycoside, which is not toxic for plants (Fig 10.15) The xenobiotic transformation mechanism includes three main stages in the detoxification of herbicides, insecticides and fungicides Phase I—at this stage reactions of degradation occur and of inclusion of new functional groups in the initial structures, as a result of which the physical activity changes as well as the hydrophilic or hydrophobic properties, the mobility of substances, which reflects their ability to pass through the membrane and the localization of the xenobiotics Primary reactions are carried out with a very high speed As a result of these reactions modified molecules have one or more groups with high reactivity (−OH, −SH, −COOH, −sNH) Inclusion of new functional groups allows further conjugation of the formed metabolites with other endogenous substances from plants Phase II (conjugation)—primary products are subjected to glycosidation, esterification, conjugation with amino acids etc The conjugation reactions most frequently involve glucose Reactions are produced by energy consumption in the presence of uridine diphosphoglucose (UDPglucose), under the action of glycosyltransferases Depending on the nature of the groups which reacted, O-glycosides, N-glycosides, and S-glycosides result Typically, glycosides and in particular the O-glycosides are highly labile substances in vitro In this context, their prolonged storage in plants would be impossible without localizing them in cell compartments, where they are protected from the action of hydrolytic enzymes At the same time, by compartmentalizing the conjugates of the xenobiotics, their action on the metabolic processes of the cell is 10.9 Biochemical Mechanism of Pollutant Transformation in Plants 303 blocked For example: through glycosidation, water solubility of conjugated substances increases, which allows vacuolar storage of the formed glycosides in the vacuolar sap Phase III—in some cases polymerization or other metabolite alterations that lead to toxin inactivation by formation of insoluble compounds The result of xenobiotic metabolization may be the detoxification (inactivation) or, on the contrary, an increase in the toxicity (activation) Therefore, while some components are detoxified, others, such as parathion, malathion, dypterex, are converted during metabolism, into even more toxic compounds (paraoxon, malaoxon, dichlorophosphonomethyl) Activation and deactivation can alternate for the same substance resulting in parallel in highly toxic compounds and in products with low toxicity Based on the above mentioned, we can generalize the biohygienic function of the plant consists not only in absorbing chemical compounds, but also in including them in the metabolism and in inactivating them Thus, plants oxidize carbohydrates forming organic acids and amino acids One of the end-products of alkane oxidation, alcohols, aldehydes, phenols, ketones is CO2, which can be used later in the process of photosynthesis by plants 10.10 Self-regulation of Plant Growth and Development in Unfavorable Environmental Conditions The reaction of various species, varieties and hybrids to unfavorable environmental conditions, quantitatively, depends on the norm of reaction, on the genotype and the capacity of self control that underlies the adaptation mechanisms of the body Comparative studies of the same genotype in different environmental conditions demonstrated that qualitatively, functional changes in the plant metabolism in different species and various stress factors are similar Thus, elevated ion concentrations have been found at high concentrations of salt, at dehydration, at temperatures below °C, under conditions of hyperthermia In all these conditions a decrease in the water content and an increase in the fluid and osmotic potential of the cell have been noticed but also changes in the functional activity of the DNA as a key element in synthetic reactions Under stress conditions there were stated disorders in the bioenergetic processes as well as in the active centers of the photosynthetic systems responsible for the primary process of transformation of the solar energy into chemical energy, for reducing the amount of free radicals in the cell and for partial blocking of the ETC This leads to a reduction of energy production efficiency as a result of photophosphorylation and oxidative phosphorylation Thus, under stress conditions, considerable amounts of energy are spent to restore the functionality and repair the damaged cell structures Simultaneously, structural changes and loss of cellular membrane integrity occurs, as a result of lipid complex oxidation as well as the deregulation of the 304 10 Physiology of Plant Resistance to Unfavorable Environmental Factors ability of metabolite compartmentalization within the cell and deregulation of the entire metabolism Analysis of the physiological dynamics of the biological parameters during stress and the nature of the relationship in each metabolic chain allows distinguishing (Fig 10.16): • primary disorders caused by direct action on the cell (disruption of osmotic processes, modification of the bioenergetics, impaired structural integrity of the membranes, reduced functional activity of the nuclear DNA) All these parameters alter significantly immediately after the onset of stress factor and, if this stress acts constantly, disorders persist throughout the entire period of the action of the factor; • secondary disorders caused by a primary disruption of the metabolic functions (changes in the processes of biosynthesis—protein biosynthesis retention, an increase in the content of inhibiting phytohormones, retention of cell multiplication and elongation Primary and secondary physiological disorders lead to the modification of important physiological functions of the vegetal organism such as the intensity of nutrient absorption and utilization, the increase in the biomass, the productivity of seeds, fruits and vegetables Damaging processes and the autoregulation mechanism that leads to adaptation under stress conditions differ by their physiological essence and are separate by the time of onset and manifestation, while the whole complex of metabolic changes under extreme conditions in vegetal cells has a phase character For the first stress phase, called the irritation phase a rapid and sudden deviation is characteristic with a rapid return to the norm of several biochemical and physiological parameters This kind of effect, which appears after a few minutes of stress, was identified, for instance, in the action of a high salt concentration on the quantity of water in chloroplasts and mitochondria, a correlation being established between the amplitude, the speed of the effect and the intensity of the stress From the physiological point of view, these changes during the irritation phase are not yet disorders of the metabolic functions, but rather specific signals of the organism about the environmental deviations from the norm This phase lasts only a few tens of minutes The next phase, called the injury phase, lasts a few days During this phase there is an inhibition of the anabolic reactions that are energy dependent and a considerable increase in the catabolic and hydrolytic reactions A strong increase in the intensity of the stress factor, which surpasses by much the maximum resistance of plants creates an imbalance between the two aspects of cellular metabolism in a manner similar to a chain reaction and, eventually, the plant dies During the irritation phase both specific and non-specific changes to various stress factors can be attested, but immediately after these changes nonspecific metabolic disorders appear that have actually a primary importance and are characteristic for the adaptation phase 10.10 Self-regulation of Plant Growth and Development … 305 Fig 10.16 Adaptation to abiotic stress (Arnholdt-Schmitt 2004) Typical phenotypic and epigenetic changes at the level of the entire vegetal organism and at the cellular level in response to phosphorus deficiency (a) and at the level of the factors involved in the global regulation of the genome (b) If the intensity of the factor does not exceed the lethal threshold, over a certain time, the ratio of anabolic and catabolic products gradually returns to normal, and initiation of the process of regeneration of the disturbed intercellular structures 306 10 Physiology of Plant Resistance to Unfavorable Environmental Factors (membranes, organelles) happens, which demonstrates the connection of the autoregulation mechanisms At the basis of the adaptation process lies the reduction of catabolic and hydrolytic reactions, since under extreme conditions, the intensity of synthetic and energy forming processes maintains the same low level as in the phase of injury It is possible that namely the weakening intensity of hydrolytic and catabolic processes is the basis and essence of the adaptation phase and reflects metabolic autoregulation in extreme conditions If the action of extreme factors ends, and optimal conditions follow (rain after drought, etc.) the repair mechanisms kick in In general, they all consist in increasing the synthesis processes previously blocked by stress, in the process of accelerated renewal of cell structures and in the adjustment of various functions to the optimum level After the action of the unfavorable factor ceases, which causes partial or total damage to the organs, their rapid regeneration begins According to the depth of physiological disorders during stress, the repair can be partial or total Plant adaptability and autoregulation processes change during ontogeny The lowest degree of adaptation is seen in young plants in the juvenile stage of growth and development and then increases slowly towards the end of the vegetation period, but is reduced during the formation of the reproductive organs, changes that correlate with the nature of the modifications during ontogenesis of the physicochemical properties of the cytoplasm All the mechanisms of self-regulation and adaptation to unfavorable conditions are realized at the cellular and intercellular level At the level of the organism they are complemented by additional mechanisms that reflect the interaction between organs During stress, competition reactions between generative and vegetative organs for water and nutritive substances are increasing If the adverse circumstances act until the differentiation of the generative organs a decrease in the number of floral primordia occurs Receiving external signals about extreme conditions, the plant forms that minimum of generative organs that it is able to supply with nutrients up to ripening If the stress factors are acting after the formation of the fructification organs, the competitive relationships begin between the seed forming organs, which constrains the development of some of them At the population level another mechanism that works effectively in adverse conditions comes into play—selection The variability of the level of resistance in a population serves as a motivation for the manifestation of this mechanism Thus, only those individuals survive who have a greater genetically determined capacity for resistance Plant adaptation to extreme environmental conditions is a complex process, coordinated by the body autoregulatory system The higher the level of biological organization (cell, organism, population), the greater the number of mechanisms involved simultaneously in the adaptation process The character of adaptive-protective reactions is unique and universal for different stress factors—salts, high temperatures, low temperatures, drought etc Glossary 307 Glossary Asphyxiation under water Plant death under conditions of excessive humidity, especially during the spring Due to the high humidity and insufficiency of oxygen plants switch to anaerobic respiration Accumulation of toxic substances like ethylic alcohol leads to cell intoxication Asphyxiation under snow Plants are perishing under a thick layer of snow (typically in winter crops), in conditions of moderate winter, at temperature near °C The cause of asphyxiation are a high rate of respiration followed by the exhaustion of the organic matter The amount of sugars in the tissues decreases from 20 to 2–4 % Uprooting Movement of the tillering node to the surface of the soil caused by the pressure exerted by the ice crust on the plant organs, which results in rupturing of the root Halophytes Plants resistant to salts which either accumulate, remove, localize salts in cell compartments or which don’t accumulate salts at all Law of B D Zalenski The leaves and other plant organs located at the top of the plant are different from the leaves and organs at the bottom by a xeromorph structure and increased resistance to drought Frost resistance The ability of plants to survive low negative temperatures Frost resistance is hereditary Salt resistance The property of plants to grow on saline soils The category of agricultural plants, resistant to salinity includes sorghum, some varieties of barley and millet and sensitive to salinity—oats and corn Heat resistance The ability of plants to support dehydration and overheating Among the drought resistant plants are the succulents, among the crop species— rice, cotton, sorghum Plant resistance The ability of plants to withstand extreme action of unfavorable factors of the environment without suffering damage This property formed during evolution and is genetic determined Winter drought The exposure of trees and shrubs to the winter winds and the sun which have a drying effect Plant tissues experience a water deficit caused by water retention by the cold ground 308 10 Physiology of Plant Resistance to Unfavorable Environmental Factors References Aleksandrov VJ (1975) Kletki, makromolekuly i temperatura L., 329 p Arnholdt-Schmitt B (2004) Stress-induced cell reprogramming A role for global genome regulation? Plant Physiol 136:2579–2586 www.plantphysiol.Oig Bohnert HJ, Nelson E, Jensenay RG (1995) Adaptations to environmental stresses Plant Cell 7:1099 Deveroll BD (1980) Zashchitnye mehanizmy rasteniy M., 126 pp Genkel’ PA (1982) Fiziologiya zharo- i zasuhoustoychivosti rasteniy M., 280 pp Grodzinskij DM (1983) Nadezhnost’ rastitel’nyh sistem Kiev, 366 pp Igamberdiev AU, Hill RD (2004) Nitrate, NO and haemoglobin in plant adaptation to hypoxia on alternative to classic fermentation pathways J Exp Bot 55(408):2473–2482 Jansen MAK, van den Noort RE, Tan MYA, Prinsen E, Lagrimini LM, Thorneley RNF (2001) Phenol-oxidizing peroxidases contribute to the protection of plants from ultraviolet radiation stress Plant Physiol 126:1012–1023 Meglickij LV, Ozeretskovskaya OL (1985) Kak rasteniya zashchishchayutsya ot bolezney M., 190 pp Nikolaevskiy VS (1979) Biologicheskie svoystva gazoustoychivosti rasteniy Novosibirsk, 278 pp Polevoy VV (1982) Fitogormony L Izd Leningradskogo universiteta, 248 pp Rubin BA, Arcihovskaya EV, Aksenova VA (1975) Biohimiya i fiziologiya immuniteta rasteniy M., 320 pp Schutzendubel A, Polle A (2002) Plant responses to abiotic stresses: heavy metal-induced oxidative stress and protection by mycorrhization J Exp Bot 53(372):1351–1365 Strogonov BI (1973) Metabolizm rasteniy v usloviyah zasoleniya M., 51 pp Tsai-Hung H et al (2002) Tomato plants ectopically expressing arabidopsis CBFL show enhanced resistance to water deficit stress Plant Physiol 130:618–626 Tumanov IM (1979) Fiziologiya zakalivaniya i morozostoykosti rasteniy M., 352 pp Index A Abscisic acid (ABA), 200, 273 Absorption, 27 Absorption maxima, 84 Absorption zone, 48 Acceptor, 94 Acetyl coenzyme A, 135 Acid gases, 293 Acropetal, 226 Action spectrum, 85 Active transport, 29, 180 Acyclic photophosphorylation, 99 Adaptation, 274 Adaptation process, 306 Adhesion forces, 58 Adsorption, 179 Aerobic dehydrogenases, 128 Aerobic respiration, 125 Aeroponics, 154 Aggregation states, 43 Agro-chemistry, Agronomic resistance, 274 Alcoholic fermentation, 132 Allelopathy, 266 Aluminum, 177 Amides, 158 Amino acid, 162 Aminotransferases, 161 Ammonia, 298 Ammonium, 157 Amorphous complex, 20 Amphiphilic, 24 Amphystomatic, 52 Amplitude, 240 Amyloplasts, 76 Anabolic reactions, 304 Anaerobic dehydrogenases, 128 Anaerobic respiration, 126 Anatomical/morphological, Annual autumn plants, 219 Annual rings, 236 Anthesis, 199 Anthocyanins, 71 Antioxidant, 67 Antiport, 30 Antisense strategies, Apical meristems, 193 Apocrine, 254 Apoplast, 20, 181, 258 Apoptosis, 125, 216 Aquaporins, 41 Arabic gum, 255 Ash, 153, 300 Asparagine, 158 Asphyxiation under snow, 276 Asphyxiation under water, 277 Assimilation factor, 102 Assimilatory cells, 70 ATP, 180 ATPase complex, 73 Autoregulation processes, 306 Auxiliary pigments, 85 Auxins, 200 B Bach, 130 Bacteriological purification, 300 Bacteriorrhizae, 155 Basipetal, 226 Benson-Calvin cycle (C3), the, 101 Biennial plants, 219 Biocenosis, 249 Biochemistry, Biological clock, 234 Biological indicators, 301 Biological repose, 192 Biological resistance, 274 Biophysics, © Springer International Publishing Switzerland 2015 M Duca, Plant Physiology, Biological and Medical Physics, Biomedical Engineering, DOI 10.1007/978-3-319-17909-4 309 310 Biopotentials, 227 Biorhythms, 226, 233 Biosphere, 10 Birch juice, 265 Blackman phase, 86 Boron, 177 Botany, Bound water, 43 Brassinosteroids, 200 Brilliant phenomenon, 119 C Cab genes, 114 Calciphiles, 172 Calciphobes, 172 Calcium, 172 Calmodulin, 173 Cambium, 189, 193 Capillary water, 46 Carbohydrates, 68, 127 Carbonate, 286 Carbon monoxide, 298 Carboxylation, 108 Carotenoids, 71 Catabolic and hydrolytic reactions, 304 Catabolism, 125 Catalase, 129 Cell death (apoptosis), 15 Cell theory, 16 Cellular, 10 Cellulose, 19 Cell wall, 16 Cerides, 256 Chelates, 152 Chelating agents, 295 Chemical and electrochemical gradients, 28 Chemical cleaning, 300 Chemical method, 154 Chemoautotrophic, 119 Chemotropisms, 223 Chinones, 94 Chloride, 286 Chlorine, 300 Chlorophyll, 68 Chlorophyll “a”, “b”, “c” and “d”, 78 Chlorophyllin, 78 Chloroplast and nuclear genomes, 110 Chloroplasts, 72 Chlorosis, 176, 301 Cholesterol, 24 Chromatoplasma, 119 Chromoplasts, 76 Chronon hypothesis, 234 Circadian rhythms, 234 Index Circalunar rhythms, 234 Circannual rhythms, 234 Circaseptal rhythms, 234 Citric acid, 135 Coenzyme Q, 138 Cohesion forces, 58 Colloidal and capillary effects, 34 Colloid hydration, 291 Colloids, 44 Compartmentalization, 291 Conducting vessels, 181 Conjugated double bonds, 90 Conjugation reactions, 301 Constitutional water, 45 Copper, 177 Copper proteids, 93 Cotransport, 29 Crassulacean acid metabolism, 101 Crinohalophytes, 288 Critical stage, 282 Crude sap, 266 Cryptochromes, 243 Crystallization water, 45 Cuticle, 51, 256 Cuticular pores, 256 Cuticular transpiration, 264 Cutin, 256 Cysteine, 165 Cytochrome, 73 Cytochrome-oxidase, 129 Cytogenesis, 190 Cytokinins, 200 Cytology, Cytophysiology, Cytoplasmic male sterility, 207 Cytoplasmic membrane (plasmalemma), 23 Cytorrhysis, 283 D Dark phase of photosynthesis, 87 Dehydrogenases, 131 Deplasmolysis, 34 Detoxification, 302 Development, 190 Dichotomyic path, 132 Diffused light, 84 Diffusion, 179, 252 Digestive glands, 255 Direct sugar oxidation, 140 Disulfide bridges, 165 Division zone, 155 Dominant centers, 196 Donor, 95 Dust, 293 Index Dwarfism, 209 Dynamic equilibrium, E Ecology, Ectodesmata, 256 Electrochemical potential, 98, 172, 180 Electron, 95 Electronic microscope, 11 Electron spins, 90 Electron transport chain (ETC), 88, 129 Electron tunneling effect, 93 Electroosmosis, 31 Electrophysiological control, 226 Electrotropisms, 223 Elongation region, 155 Embryonic stage, 195 Emerson effect, 87 Endogenous, 45 Endogenous duration, 240 Endoplasmic reticulum, 24 Endosmosis, 155 Energetic level, 89 Energogenesis, 126 Energy and matter circulation, 69 Environmental factors, 10 Enzymes, 17 Epinasties, 224 Erythrose, 141 Esterification, 302 Ethylene, 200 Etiolation, Etioplasts, 76 Euhalophytes, 288 Exchange of ions, 179 Excitation state, 90 Excretion, 27, 250 Exergonic, 138 Exogenous, 45 Exometabolites, 250 Experimental science, Extrafloral nectaries, 258 Exudates, 253 F Facilitated diffusion, 29, 180 Facultative halophytes, 289 Fatty acid synthesis, 138 Fermentation, 125 Ferredoxin, 94 Ferritin, 176 Ferroprotein, 138 Fe-S proteins, 92 Field experiments, 154 311 Flavoproteins, 93, 128 Floral clock, 237 Fluctuation period, the, 240 Fluorescence, 90 Fluorine, 300 Forced dormancy, 192 Free water, 43 Frost resistance, 279 Fructose-6-phosphate, 145 Fumaric acid, 135 G Gametophyte, 197 Gas exchange, 126 Gelation, 257 Genetically modified plants (GMPs), Genetic program, 199 Genetics, Geotropisms, 223 Gibberellins, 200 Glandular hairs, 259 Global warming, 119 Glucose, 135 Glucose-6-phosphate, 144 β-glucuronidase, 233 Glutamate dehydrogenase, 159 Glutamine, 158 Glutathione, 166, 298 Glycohalophytes, 288 Glycolic acid, 109 Glycolic pathway of carbon transformation., 107 Glycolipids, 24 Glycolysis, 132 Glycophytes, 287 Glycosidation, 302 Glyoxylate cycle, 140 Golgi apparatus, 24 Granal and stromal, 73 Gravitational water, 46 Growth, 189 Growth and development, Growth inhibitors, 201 Growth promoters, 201 Growth regulators, Growth respiration, 126 Guard cells, 51, 171 Guttation, 46, 58, 263 H Halophytes, 287 Hardening, 278 Hatch-Slack-Karpilov cycle (C4), 101 Heat-loving plants, 277 312 Heat shock proteins, 273 Heavy metals, 294 Hecht’s filaments, 34 Heliophytes, 47, 79 Hemicellulose, 19 Herbicide, 302 Hexochinase, 144 Hexoses, 132 Hill phase, 86 Hipostomatic, 52 Holocrine, 254 Homeohydrophytes, 60 Homeostasis, 267 Hyaloplasma, 21 Hydathodes, 257 Hydatophytes, 60 Hydration water, 280 Hydroactive, 53 Hydrogen sulfide, 298 Hydropassive, 53 Hydrophilic, 24 Hydrophobic, 24 Hydroponics, 154 Hydrotropism, 48, 223 Hygrophytes, 60 Hygroscopic water, 45 Hyperstomatic, 52 Hypertonic, 33 Hyponasties, 224 Hypothesis of the multioscillatory model of biorhythms, 236 Hypotonic, 34 Hystogenesis, 190 I Idioblasts, 259 Imbibition, 31 Imbibition force, 44, 54 Inanition, 276 Inductive resonance, 91 Infrared, 82 Injury phase, 304 Inorganic ions, 17 Insecticides, 301 Insectivorous plants, 248 Insoluble compounds, 303 Intensity of respiration, 128 Intercalary meristems, 193 Intercellular, 10 Intrafloral nectaries, 258 Ionic asymmetry, 172 Ion pumps, 29 Irrigation, 285 Irritation phase, 304 Index Isocitrate dehydrogenase, 145 Isocitric acid, 135 Isotonic, 33 J Juvenile stage, 196 K Kaspari stripes, 48 α-ketoglutarate dehydrogenase, 145 Ketoglutaric acid, 135 Kinases, 112, 168 Krebs cycle, 126 L Lactiferous cells, 259 Latency, 189 Lateral meristems, 193 Latex, 255 Law of Müller and Hegel, 61 Leghemoglobin, 164 Leguminous plants, 162 Lenticels, 52 Lenticular transpiration, 264 Leucoplasts, 76 Light-harvesting complex, 73 Light phase of photosynthesis, 87 Light quantum, 82 Light utilization coefficient, 83 Lignin, 254 Liposolubility, 180 Localization signal, 75 Long day plants, 218, 238 Lysosomes, 24 M Macroelements, 154 Macroergic bonds, 126 Macroergic ester bonds, 169 Magnesium, 174 Maintenance respiration, 126 Malate dehydrogenase, 145 Malic or oxaloacetic acids, 107, 135 Manganese, 175 Mathematical modeling, 11 Maturation, 216 Mature region, 155 Mechanical cleaning, 300 Median lamella, 267 Membrane hypothesis, 236 Meristematic tissues, 189 Merocrine, 254 Mesophyll cells, 118 Mesophytes, 60 Index Metabolic disorders, 304 Metabolic water, 282 Metabolite compartmentalization, 304 Metal oxides, 293 Metamers, 236 Methane, 298 Methionine, 165 Micro-and macrofibril complex, 20 Microelements, 154 Middle lamella, 18 Mineral elements, 152 Mineralization, 152, 257 Mineral nutrition, Mitochondria, 125 Mitochondrial cristae, 129 Mn-Mn dimers, 97 Molecular, 10 Molecular oscillation, 241 Molybdenum, 176 Morphogenesis, 6, 190 Mucilage, 257 Multicellular, 16 Mycorrhizae, 155 N Na+/K+pumps, 31 Na/K ratio, 290 NADPH+H+, 87 Nasties, 222 Necrosis, 166, 301 Nectar, 250 Nectary glands (nectaries), 255 Neutral plants, 218 Nitrate reductase, 158 Nitrates, 157 Nitrites, 157 Nitrogen, 156 Nitrogenase, 164 Nitrogen oxides, 300 Nitrogenous bases, 162 Nutations, 223 Nyctinastic movements, 225 O Obligatory halophytes, 289 Ontogenesis, Ontogenetic dynamics, Organelles, 21 Organic acids, 288 Organismal, 10 Organogenesis, 190 Organogenic elements, 152 Orthophosphoric acid, 168 Osmophores, 255 313 Osmosis, 31, 42 Osmotic force, 44, 54 Osmotic pressure, 32, 286 Osteole, 52 Oxidases, 129 Oxidation cycle, 130 Oxidative degradation, 67, 125 Oxysomes, 138 P P680, 89 P700, 89 Pacemaker, 236 Palisade parenchyma, 70 Palladin, 130 Passage cells, 48 Passive transport, 28, 179 Pectic substances, 19, 173, 257 Pedological and atmospheric drought, 284 Pedology, Pellicular water, 45 Pentanonic homocycle, 79 Pentose phosphate cycle, 140 Perennials, 219 Pericycle, 193 Perivascular sheath cells, 109 Permanent wilting, 280 Peroxidase, 129 Peroxisomes, 108 Pesticides, 301 Phellogen, 189, 193 Phenolic compounds, 253 Pheophytin, 92 Phosphatases, 112 Phosphoenolpyruvate, 145 Phosphoenolpyruvic acid, 103 Phosphofructokinase, 144 3-phosphoglyceric acid, 102 Phosphoglyceric acid, 133 Phosphoglyceric aldehyde, 103, 133 Phospholipid molecule, 168 Phospholipids, 24 Phosphorescence, 90 Phosphorylation, 168 Photoactive, 53 Photoelectric effect theory, 86 Photoinactivation, 118 Photons, 82 Photooxidation, 70 Photooxidative stress, 72 Photoperiodism, 218, 236 Photoprotection, 71 Photorespiration, 101 Photosynthesis, 314 Photosynthetic apparatus, 111 Photosystems I (PSI) and II (PSII), 73 Phototrophic, 119 Phototropisms, 223 Phragmoplast, 254 Phycobilins, 77 Phycocyanin, 80 Phycoerythrin, 80 Phyloquinone, 92 Physiological, Physiological drought, 286 Physiologically acid soil, 183 Physiologically basic soil, 183 Physiological method, 154 Phytin, 169 Phytochrome, 80, 222, 232 Phytohormone regulation, 226 Phytohormones, 9, 17, 200 Phytol, 78 Phytoncides, 300 Phytopathology, Phytotoxicity, 299 Phytotrons, 10 PIF3, 243 Pinocytosis, 31 Plant breeding, Plant cultivation, Plant morphology, Plant productivity, Plasmodesmata, 20, 256 Plasmolysis, 33 Plastids, 16 Plastochron, 236 Plastocyanin, 92 Plastoquinone, 92 Poichilohydrophytes, 60 Polarity, 226 Polyphenols, 129 Population, 10 Porphyrin core, 79, 129 Potassium or kalium, 170 Primary acceptor, 92, 94 Primary and secondary metabolites, 151 Primary cell wall, 18 Primary disorders, 304 Procambium, 193 Processing, 113 Progressive branch of nitrogen exchange, 162 Proplastids, 75 Protein carriers, 29 Protochlorophyllides, 117 Protoplasm, 23 Protoplasm colloidal status, 152 Protoplast, 19 Index PRR, 244 Pyrophosphate bonds, 169 Pyrrol, 79, 129 Pyruvic acid, 132 Q Quantasomes, 73 Quinones, 129 R Radial and xylemic transport, 178 Radial transport, 55 Radiation, 295 Radicals, 301 Radioprotective substances, 298 Radiosensitivity, 296 Reaction centre, 88 Redox reactions, 79, 131 Redox system, 176 Regressive branch of nitrogen exchange, 160 Regulatory systems, Relative transpiration, 50 Reproductive development, 190 Reproductive stage, 196 Resistance, 273 Resistance to drought, 280 Respiration, Respiratory coefficient, 127 Respiratory pigments, 131 Respiratory substrate, 127 Retardants, 201 Rhizobium genus, 162 Rhizosphere, 48, 250 Ribulose-1,5-bisphosphate carboxylase (RUBISCO), 74 Root cap (calyptra), 155 Root hair region (differentiation zone), 155 Root hairs, 48 Rooting, 206 Root nodules, 162 Root pressure, 56 S Saline soils, 166 Salinization, 286 Salt glands, 291 Sciophyts, 47, 79 Secondary acceptor, 95 Secondary cell wall, 18 Secondary disorders, 304 Secretion, 27, 250 Sedoheptulose, 141 Seismonastic movements, 225 Seismotropisms, 223 Index Self-regulation (autoregulation), Semipermeability, 21 Senescence phase, 196 Short-day plants, 218, 239 Silicon, 176 Simple diffusion, 28 Singlet, 90 Soil colloids, 166 Soil solution, 182 Spectral composition, 116 Spongy parenchyma, 70 Sporophyte, 197 Stomata, 51 Stress conditions, 292 Stress metabolites, 250 Stroma, 75 Structural molecules, 17 Suber, 254 Suberization, 257 Succinate dehydrogenase, 145 Succinic acid, 135 Succulence, 290 Suction force, 31, 55 Sugars, 288 Sulfate, 165 Sulfur, 165 Sulphate, 286 Symplast, 20, 181, 258 Symport, 30 T Tearing, 263 Temporary wilting, 280 Termonastic movements, 225 Terpenes, 250, 259, 267 Thermoperiodicity, 245 Thioredoxin, 117 Thylakoids, 73 Tonoplast, 23 Tonoplast transferases, 295 Toxic gases, 298 Tracheids, 51 Transamination, 160 Transcription, 114 Transition to flowering, 199 Translation, 113 Transphosphorylation, 169 Transpiration, 263 Transpiration coefficient, the, 50 Transpiration intensity, 50 315 Transpiration productivity, 50 Transpiration pull, 57 Tricarboxylic acid cycle, 135 Triplet, 90 Trophic regulation, 226 Trophic substances, 266 Tropism, 222 Turgidity, 33, 43, 171 Turgor pressure, 35 U Ubiquinone, 92 Ultracentrifugation, 11 Ultramicroelements, 154 Ultrastructure, 11 Ultraviolet, 82 Uprooting, 277 Upward flow of water, 286 V Vacuolar Na +/H + antiporters, 292 Vacuole, 16 Vegetative development, 190 Vapors of acids, 293 Vernalization (yarovization), 218 Visible spectrum, 81 W Water aggregation states, 43 Water balance, 58 Water oxidation, 97 Water regime, Waxes, 250 Wilting coefficient, 283 Wilting point, 46 Winter drought, 307 X Xanthophyll, 89 Xenobiotics, 300 Xeromorphic features, 61 Xeromorphic qualities, 284 Xerophytes, 60 Xylem vessels, 57 Z Z-scheme, 94 Zeatin, 211 Zinc, 177