Chapter 1

Introduction to the Educational

Course of Plant Physiology

Abstract Plant physiology is a science that studies vegetal organisms in ontoge-

netic dynamics—the diversity, the laws and the mechanisms of physiological and
biochemical processes, their biological significance, their dependence on environ-
mental factors. Traditionally, it was based on two directions: anatomical/morpho-
logical and physiological, but this division is somewhat relative, because structure
and function have evolved in parallel and cannot be studied separately. This
interdisciplinary research field focuses on a series of compartments like: plant cell
physiology; water regime; photosynthesis; mineral nutrition; respiration; growth
and development; resistance to unfavorable factors; phenomena of self-regulation at
all the levels of organization (including at the organism level by means of inter-
acting centers of dominance). While as a theoretical science plant physiology tries
to obtain an integrated, detailed picture of the molecular, biochemical, physiolog-
ical, morphogenetic processes going on in the living plant and the interconnection
between these, at the applicative level its aim is to be able to direct vital processes in
the life cycle of a plant like growth, development, metabolism, photosynthesis,
nutrition, resistance, fructification in order to control the vitality or yield of the crop
species and to maximize economic benefits. Classical research in plant physiology
is carried out in the field, in vegetation pots, solariums, greenhouses, phytotrons,
laboratories. Experiments make use of a diverse range of methods like: imaging
technologies (optical and electronic microscopy), centrifugation, chemical analysis,
chromatography, radioactive labeling, gel filtration, electrophoresis, X-ray analysis,
in vitro culture, but also in silico mathematical modeling to predict the behavior of
various systems and the output parameters.

© Springer International Publishing Switzerland 2015 1

M. Duca, Plant Physiology, Biological and Medical Physics,
Biomedical Engineering, DOI 10.1007/978-3-319-17909-4_1
2 1 Introduction to the Educational Course of Plant Physiology

Historical Background
1727—St. Hales identifies the pathways of water, mineral salts and organic sub-
stances circulation.
1771—J. Priestley discovers photosynthesis.
1775—M. Malphigi describes the cycle of substances in plants—the ascending and
descending currents.
1800—J. Senebier edits “Plant Physiology” in 5 volumes.
1804—J. Senebier and Th. Saussure argue that photosynthesis represents the
nutrition of plants with carbon.

Brief Updates
During the last decades, by using gene engineering methods, plants with recom-
binant DNA have been created, also called genetically modified plants (GMPs), this
fact favoring the emergence of a new direction in plant physiology—the physiology
of transgenic plants which aims to determine the physiological and biochemical
changes of transgenic plants as a result of the inclusion of new genes into their
genome. Thus, the use of GMPs has allowed the elucidation of the genetic and
physiologic mechanisms of the activity of genes artificially included in the plant
1 Introduction to the Educational Course of Plant Physiology 3

genome, among which are also those that are normally found in animal organisms,
such as the Green Fluorescence Protein gene (GFP) from certain jellyfish species.
The GFP emits a green fluorescence under UV light, and its fusion with any other
protein allows the positional analysis of the last within the cell, the mechanism
being similar to that of radio-labeling.
Inserting auxine phytohormone biosynthesis genes (iaaM and iaaH) into the
tobacco genome resulted in more viable transgenic plants with a more active
vegetative morphogenesis and reproductive development and with both a higher
amount of water stored in tissues and a higher resistance to drought.
Another example is represented by the ferric superoxide dismutase gene
(FeSOD) from Arabidopsis thaliana (one of the genes involved in antioxidative
protection) which was included into the genomes of tobacco and wheat. The
genetically modified plants proved more resistant to the oxidative stress than the
control, confirming that the gene is expressed.
Lately, to study a particular gene function the antisense strategies are often
applied. The best known example is given by the gene that encodes the synthesis of
the polygalacturonase enzyme, involved in cell wall degradation in ripening tomato
fruits. After including this gene in the tomato genome, in reverse orientation, sense
and antisense RNA will bind on the basis of complementarity, thus obstructing
translation and leading to longer fruit preservation.

1.1 The Definition and Scope of Plant Physiology

Plant physiology is a very important branch of biological sciences that studies the
life of plants—the laws and mechanisms of physiological and biochemical pro-
cesses, their significance, their interdependence with environmental factors in
ontogenetic dynamics. The notion of physiology originated from Greek by joining
the words physis, which means “function” and logos—“science”.
Plant physiology has appeared in 1800, when the Frenchman J. Senebier edited
his first monograph in five volumes “Plant Physiology”, which included not only
his own experimental results, but also those obtained in this scientific field by:
M. Malpighi, who has described the flow of substances in the plant (1775);
St. Hales, who demonstrated that water and mineral salts flow through the xylem,
while organic substances—through the phloem (1727); J. Pristley, who has dis-
covered photosynthesis (1771), etc.
During the development of plant physiology as a science, it has been based on
two directions: anatomical/morphological (descriptive) and physiological (experi-
mental), which, in principle, can be considered two basic research methods. This
division is relative, because vegetal organs can’t be studied without taking into
account their function, just as any processes cannot be studied without knowing the
4 1 Introduction to the Educational Course of Plant Physiology

structures they are localized in. Any physiological process should be regarded as a
product of long evolution, which forms the plant ability to adapt to variable
environmental conditions. The function has evolved in relationship with the
structure of the organism and the structure has stabilized under the action of
environmental factors and according to the function. Thus, to study respiration, it is
necessary to know the structure and ultrastructure of mitochondria, and to reveal the
mysteries of photosynthesis, a unique and specific process happening only in green
plants, it is important to know the structure and ultrastructure of the assimilatory
apparatus.
Most of the compartments of plant physiology have been delimited in the
nineteenth century and are valid even nowadays. These are:
1. Cytophysiology (plant cell physiology);
2. Water regime of plants (H. Dutrochet, H. de Friz, J. Sachs);
3. Photosynthesis (G. Busengo, M. Ţsvet, J. Pristley, K.A. Timireazev);
4. Mineral nutrition (I. Leibih, G.B. Busengo, D.N. Preanishnikov);
5. Respiration (A.S. Famiţsin, I.P. Borodin, L. Paster);
6. Growth and development (J. Sachs, A.S. Famiţsin);
7. Plant movements (T. Nait, J. Sachs, Ch. Darwin);
8. Irritation (B. Sanderson, Ch. Darwin);
9. Resistance to unfavorable factors (D.I. Ivanovski).
Thus, plant physiology as a distinct branch of biology, aims to study successively
all vital processes that occur in vegetal organisms. In the second half of the twentieth
century the basics of a new branch of plant physiology named self-regulation were
laid. The phenomena of self-regulation and coordination of physiological processes,
as well as other processes, are studied at all the levels of organization of living matter
(molecular, intracellular, at the levels of tissue, organ, organism, biocoenosis) the
mechanisms of implementation being diverse and specific.
Self-regulation (autoregulation) is the property of biological systems to maintain
the stability of the physical and chemical conditions of the internal environment, of
the structure and properties of the organism in their elementary form, all these in
conditions of a dynamic equilibrium. Autoregulation represents the process, which
minimizes various deviations in the biological systems (pH, viscosity, redox-
potential, etc.), resulting from the influence of causative agents. Therefore, the
capacity of the vegetal organism of carrying out vital functions amidst changing and
unfavorable environmental conditions is implemented.
Such a stability has a dynamic and active character. It is maintained by complex
mechanisms, which determine the coordinated physiological activity of different
organs, thus allowing autoregulation of plant growth and development, organism
temperature, raw sap composition, regeneration of damaged tissues, adaptation to
stress conditions, etc. (Figure 1.1).
Self-regulation ensures integrity and homeostasis of plant organisms, allows
harmonious growth and development and helps react adequately to the alternating
1.1 The Definition and Scope of Plant Physiology 5

Fig. 1.1 Scheme representing regulative and directive processes in living organisms (Polevoy
1989)

factors of the environment. Autoregulation mechanisms are turned on automatically

in the appropriate place and time, according to the needs of the organism.
The notion of self-regulation is characteristic both for the whole vegetal
organism and for the individual cells. In fact, it is at the cellular level that the
integration of plant physiology with genetics, cytology, molecular biology, bio-
chemistry, biophysics, etc. happens.
Study of the autoregulation phenomena may contribute to the transition from
describing the processes happening in plant organisms to their direct manipulation
by acting upon the corresponding regulatory systems. In the last decades, the number
of theoretical and experimental studies dealing with regulation and autoregulation of
gene and enzyme functional activity, with membrane, electro-physiological,
phytohormonal control (particularly related to the development of gene engineering
and biotechnology) has considerably increased. Self-regulation determines plant
homeostasis and creates conditions for the epigenesis of functions, which implies a
strong collaboration between the factors of the environment and the plant genome.
Consequently, it leads to the appropriate phenotypic expression.
Regulatory systems (Fig. 1.2) at the cellular level include:
• the mechanisms that determine qualitatively the enzymatic equipment of the
cells and which consequently determine the metabolic profile of the cells, tissues
and organisms;
• the mechanisms maintaining a relative constant of the cellular metabolism
(quantitative regulation of enzyme activity, of membrane transport etc.).
All these regulatory systems are interdependent. For example, gene activity
determines the properties of the cell membrane, and biological membranes also
influence the differential activity of the genes.
With the advent of multicellular organisms, intercellular regulatory systems have
emerged including trophic, hormonal, electrophysiological regulation, contributing
6 1 Introduction to the Educational Course of Plant Physiology

Dominant centers

Polarity Biorhythms Channel connections

Regulatory contours

Electrophysiological
Phyto hormonal regulation
regulation

Trophic regulation

Genetic regulation Membrane regulation

Enzymatic activity regulation

Fig. 1.2 Interaction of regulatory systems (Polevoy 1989). Regulatory level: I intracellular, II
intercellular, III organismal

to the interaction between plant organs. Such an interaction can be observed during
cultivation of different vegetative explants in vitro. However, the existence of
trophic, hormonal and electrophysiological interactions between cells, tissues and
organs does not fully explain the behavior of a plant as a whole living organism.
There are higher level regulatory systems and mechanisms connecting organs and
functional systems of the plant during the life cycle of the organism and its onto-
genetic transitions.
The basic autoregulation mechanism at the organism level relies on the presence
of a few centers of dominance (the stem and the root apexes), which receive
information both from the external and internal medium and influence the living
organism by driving tissue morphogenesis, by creating physiological gradients,
polarity, channel connections (conductive fascicules), physiological oscillations.
1.1 The Definition and Scope of Plant Physiology 7

Polarity and channel connections coordinate the space orientation of morphological

processes, while oscillations help achieve time coordination.
At the organism level these regulatory centers unite in regulatory contours with
reversible relationships—positive or negative, which constitute the irritability effect.
Thus, the integrity of the plant organism is determined by the interaction of control
systems with central directing elements represented by the dominant centers.
All the physiological processes occurring in plants are studied from various
aspects.
From the biochemical aspect, plant physiology studies the biosynthesis of
organic compounds from inorganic ones, the functional importance of the diversity
of organic substances formed as a result of the primary and secondary metabolism
of the plant. It also researches the pathways of photosynthesis, reveals the laws of
mineral nutrition, the importance of mineral substances as regulators of metabolic
processes, their role in electrical phenomena occurring inside the cell or in the
synthesis of organic compounds, etc.
From the biophysical aspect the following problems are studied: cell energetics,
electrophysiology of vegetal organs, physical and chemical laws of the water
regime, those of nutrition via the root system, of growth, photosynthesis, respira-
tion, the electrical aspects of irritability.
From the aspect of evolution, researchers study the physiology of the genus,
species, individual, as well as ontogenesis as a function of the genotype, which has
transformed during phylogenesis.
The ontogenetic aspect implies the analysis of the age-related laws governing
plant growth and development based on the physiological and biochemical processes
occurring in cells, tissues, organs as well as the study of morphogenesis and possible
ways of acting on plant development (interfering with the photoperiodism, hardening
plants, manipulating phytohormone signaling pathways to control plant stature, etc.).
The ecological aspect consists of studying the dependence of the internal
processes and of the particularities of individual development of the vegetal
organism on the multitude of environmental conditions.
Plant physiology is an experimental biological science that summarizes the
ensemble of theoretical and practical knowledge, based on which, by using the
principles of the scientific method, it is possible to intervene in the most important
processes in the life cycle of plants: growth, development, metabolism, photosyn-
thesis, nutrition, resistance, fructification.
In describing the studied phenomena, plant physiology integrates knowledge
from different areas of biology and life sciences such as: botany and plant mor-
phology (studying the structure and components of the vegetal organism), cytology
(studies the cells), biochemistry (investigates chemical substances and reactions
occurring in living organisms), biophysics (focuses on the description of physical
phenomena related to living organisms for instance energy exchange between
plants and the environment, etc.), ecology (provides data on the effect of envi-
ronmental factors on plants), chemistry, physics, mathematics, etc. At the same
time, plant physiology represents a theoretical basis for plant cultivation, phyto-
pathology, plant breeding, agriculture, agro-chemistry, genetics and pedology.
8 1 Introduction to the Educational Course of Plant Physiology

1.2 Purposes of Plant Physiology as a Science

The possibility to constantly build up on available research knowledge and the

potential for implementing the final results make plant physiology a fundamental
science with practical applicability.
The purpose of plant physiology as a science is to investigate the peculiarities of
the life of different species of vegetal organisms both cultivated ones and those from
the spontaneous flora, in order to direct key processes like growth, development,
nutrition, metabolism and others.
Being a fundamental science, plant physiology aims to examine the molecular,
physiological, biochemical and morphogenetic mechanisms of the vital processes in
their dynamical succession and as a function of alternating environmental condi-
tions, including:
• discovering the essence of the organism’s individual development and studying
the interaction of genetic, physiological, enzymatic mechanisms during growth
and development;
• elucidating regulatory and autoregulatory mechanisms under the action of
external factors;
• detailing the biochemical theory concerning mineral nutrition of plants;
• elucidating the ways used by plants to improve the efficiency of solar energy
utilization;
• investigating the mechanism of atmospheric nitrogen fixation and its utilization
by superior plants;
• developing and detailing the theoretical bases of the use of biologically active
substances;
• elucidating the laws of plant viability (mechanisms of nutrition, growth,
movement, reproduction);
• improving the theoretical knowledge on maximizing crop yields;
• researching endogenous mechanisms of regulating physiological functions,
including basic mechanisms of enzyme biosynthesis, transport of substances and
regulatory action of biomembranes;
• decoding mechanisms that control the chronological sequence of genetic pro-
gram implementation during plant ontogenesis, including intracellular interde-
pendence, interaction between vegetal organs during growth, reproduction and,
finally, during crop formation;
• studying the regulation of secondary metabolite biosynthesis (alkaloids, rubber,
phenolic compounds, etc.) which are often of great economical importance.
As an applicative science, plant physiology aims to increase plant productivity.
It is known that in an ear of wheat there are 3–5 flowers, of which only 1, 2 or 3 are
fructifying. To make all these flowers fructify is the kind of challenges put in front
of plant physiology. To achieve this goal, it is necessary to know the causes
preventing metabolite formation and grain filling. At the core of such phenomena
lies, for instance, the poor activity of the photosynthetic apparatus, caused by the
1.2 Purposes of Plant Physiology as a Science 9

depletion of chloroplast enzymes or the exhaustion of cell energy resources in the

form of adenosine triphosphate. In order to solve tasks like this, plant physiology
investigates:
• photosynthetic apparatus activity and efficiency of solar energy use;
• plant requirements for mineral nutrition;
• water regime and efficiency of water utilization;
• plant resistance to various unfavorable factors;
• the possibility of using growth regulators;
• critical phases of ontogenesis;
• physiological bases of implementing the morphogenetic program.
A problem of the physiology of mineral nutrition, with promising prospects in
plant breeding, is to study the absorption of nutrients by the root system. Knowing
the rhythm and the rate of nutrient absorption in the multitude of varieties resulting
from plant breeding, we can choose the biological material with a maximum
capacity of fertilizer intake, this being a prerequisite for big crops.
Discovering the functions of growth regulators may have multiple practical
applications. Thus, gibberellins can be used to spray tree seedlings in greenhouses
to force their growth in the first year and to reduce the overall time spent in a
greenhouse, while auxins can be used to stimulate seedling rooting. It is again the
task of plant physiology to determine experimentally for various species the
duration of seedling exposure to these phytohormones, their optimal concentrations
and the optimum age of the treated sprouts in order to achieve the best results.
Another important task of plant physiology is to find out plant requirements with
regard to nutrients and water during different vegetation periods. In autumn cereals,
regrowth of the foliar system that has been destroyed during the winter by the frost
involves the consumption of large amounts of nitrogen in the early spring.
In vegetable production, the practical purpose is to obtain seedlings in green-
houses during winter, when the light intensity is low because of permanent
cloudiness. The etiolation phenomenon (plants have long, weak stems; smaller,
sparser leaves due to longer internodes; and a pale yellow color) can be prevented
by illuminating plants with artificial light or spraying them with diluted solutions of
retardant substances that hamper seedling elongation.

1.3 Research Methods Used by Plant Physiology

As mentioned above, plant physiology is a pronouncedly experimental science, the

experiment representing the main research method. The experiment is preceded by
the hypothesis. Studies and experiments on plants are performed under three basic
aspects:
(1) Passing from a higher level to a more elementary one, from analyzing complex
biological laws to studying simpler ones—physical and chemical. This
10 1 Introduction to the Educational Course of Plant Physiology

research direction led to the advent of molecular biology, to the discovery of

the hereditary code, the protein biosynthesis mechanism, the main laws of
absorption and utilization of light quanta in photosynthesis. This, however, is
not sufficient to understand the laws of physiological processes occurring in
vegetal organisms.
(2) Currently another approach is used—from simple to complex, called by V.A.
Engelhardt integral. Generally, this path allows pursuing the evolution of
certain processes at the level of DNA–RNA–protein–enzyme–biochemical
reaction–physiological process–property of the cell. At any level of this path,
regulation is possible and there are also internal mechanisms of autoregulation
(targeting DNA replication, RNA and protein biosynthesis, enzymatic activity,
but also cell, tissue and organ differentiation).
(3) Physiological processes are studied in ontogenetic dynamics and in relation-
ship with environmental factors.
Plant physiology research is carried out in the field, in vegetation pots, solari-
ums, greenhouses and laboratories. The most modern investigations are carried
nowadays in phytotrons (the name was given by R.A. Millikan in 1949). Phytotrons
are constructions with special rooms with either natural or artificial illumination,
heated or cooled artificially, with adjustable air temperature and humidity. With the
help of automatic electrical installations, vegetation parameters can be maintained
at certain established levels. Investigations in the field of plant physiology imply
carrying out experiments and exploring processes and phenomena at different levels
of organization of the living matter by means of biochemical, biophysical, physical-
chemical and biological methods.
At the molecular level the physical-chemical processes occurring in living
organisms are studied: the synthesis, assembly and restructuring of the proteins,
nucleic acids, polysaccharides, lipids and other substances, the energetic and
informational metabolism that regulate these processes. At the cellular level the
structure and properties of the cell and its components is studied, the relationship
between organelles etc. At the intercellular level knowledge from multiple disci-
plines is integrated and the principles of photosynthesis, respiration and interaction
between tissues and organs are provided. At the organismal level the processes and
phenomena taking place in an individual organism are studied as well as the
coordinated functioning of its organs and systems, interactions between different
organs and their individual roles, changes caused by accommodation.
At the population level research focuses on the basic unit of the evolutionary
process—the population. It means that the interactions between individuals that
inhabit a certain territory (more or less isolated) is investigated. The composition
and dynamics of the population is strongly correlated with the molecular, cellular,
intercellular and organismal levels of organization. At the level of the biosphere the
processes taking place in the biogeocenoses are studied including the interactions of
biotic and abiotic components of the ecosystems.
Each of the mentioned levels of organization has its own specific research
methods. The observation of various phenomena is carried out with the naked eye
1.3 Research Methods Used by Plant Physiology 11

(macroscopically) or using a microscope (microscopically). The discovery and

constant improvement of imaging technologies with the electronic microscope had
marked a significant stage in plant physiology development—the era of cellular
organelle physiology. The electronic microscope together with the range of meth-
ods for cellular homogenate ultracentrifugation allowed the study of the submi-
croscopic structure of cellular organelles, while chemical micro-analysis allowed
elucidation of their chemical composition. Based on knowledge about the ultra-
structure and chemical composition it has become possible to decode physiological
functions of various cell organelles.
The scientific works in the field of plant physiology often make use of: ordinary
microscopy, electronic microscopy, centrifugation, chemical analysis, chromatog-
raphy, radioactive labeling, gel filtration, electrophoresis, roentgen analysis, artifi-
cial modeling of systems, autoradiography, in vitro culture.
Lately, side by side with physiological and biochemical methods, mathematical
modeling of life processes and plant productivity in defined conditions of growth
and development is used by utilizing model-algorithm-program triads.

Glossary

Adaptation The evolutionary process by which the organism or species survives

and reproduces in new environmental conditions.
Self-regulation (autoregulation) The general feature of biological systems that
assures the control and autonomous coordination of the functioning of system
elements and the maintenance of a dynamic equilibrium in the system.
Evolution The progressive development of living organisms during successive
generations by means of accumulating favorable hereditary variations enforced
by natural selection.
Enzyme A protein produced by the cell which controls the reactions of synthesis
and degradation via its catalytic activity, playing a fundamental role in metabolic
processes regulation.
Phylogenesis The history of the development of a species or other taxonomical
unit during the evolutionary process.
Phytohormone A substance secreted by the plant cell in small quantities, which
controls various aspects of growth, developmental transitions, organ morpho-
genesis, response to various stress factors etc.
Photosynthesis The fundamental process of synthesis of organic compounds from
inorganic ones (CO2 and H2O) in the presence of light, carried out by green
plants and photosynthesizing microorganisms. During the process of photo-
synthesis the solar energy is transformed into the energy of chemical bonds in
organic molecules.
12 1 Introduction to the Educational Course of Plant Physiology

Metabolism The totality of all the complex processes of synthesis (energy storage)
and degradation (energy release) undergone by the substances in a living
organism.
Morphogenesis Cyto-differentiation and development of visible structures (organs
or parts) in an organism during ontogenesis.
Levels of organization Systems with a specific organization (characteristic of
biological systems only) and with a character of universality.
Ontogenesis The series of transformations undergone by the organism, from egg
fecundation to death, according to the scenario for the respective species.
Respiration The process of oxidative degradation of complex organic substances
into inorganic ones accompanied by energy release.

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