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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.
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)
Dominant centers
Regulatory contours
Electrophysiological
Phyto hormonal regulation
regulation
Trophic 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
Glossary
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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