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Lecture On Plant Physiology
Lecture On Plant Physiology
Sand culture technique is also employed. Except for Hydroponics have certain advantages like:
possibility to provide desirable raising crops in green houses, these techniques are not
economically feasible at present, nutrient environment; regulation of pH of the nutrient solution
conducive to specific crops; it is possible to replace the nutrient solution periodically; it is
possible to control pathogens which are otherwise present in the soil; it is feasible to regulate
aeration at regular intervals: in situations where equipment is automatic much cost on labour can
be saved; there is no need to undertake tillering weeding; uniform plant growth can be easily
achieved and nutrition budget can be easily regulated to meet the changing pattern of plant
growth and maturity.
However, hydroponics have some disadvantages also and these are: limited plant yield and
production; the system is only suitable to more specific and highly valued crop species;
considerable technical experience and skill is needed to handle hydroponics; each crop demands
specific modifications, and lastly several of the crop plants like potatoes, sweet potatoes, carrots
may even change their shape and size.
Despite its several disadvantages and practical difficulties, large scale commercial hydroponics
growth is carried out in many floricultural and horticultural species. In several of the European
countries it has been possible to obtain high yields in radish, tomato, lettuce, cucumbers, etc.
Pectic compounds and hemicellulose play an important role in the initial stage of cell growth.
As the cell matures secondary wall starts developing on the inner surface of the primary wall.
The cell wall thickens as layers of cellulose are laid down by the cytoplasm.
The wall becomes plastic and the cell growth ceases. Secondary wall gives the plant cell its
structural independence and mostly contains cellulose and other polysaccharides including
hemicellulose.
It is less hydrated and hence more dense. The secondary wall is mostly mechanical in function
and is usually distinguishable in three layers (S1, S2, and S3).
Of these, the S3 layer is also referred to as the tertiary wall. In some plant species the secondary
wall is made up of several layers which are designated as S1, S2 … Sn etc. It is a permanent
layer i.e. it cannot be extended by growth.
Cytoplasmic ground substance (cytosol) is not a constant material and its consistency changes
from time to time.
Its main component is water and its percentage varies from 90 (parenchyma cells) to 4-5 (in
seeds; spores and pollen).
The cytosol contains several living (mitochondria, plastids, ribosomes, lysosomes, etc.) and non-
living particles (fat globules, aleurone grains, organic acids, pigments, fat globules, etc.).
Cytosol varies enormously in the same cell at different times and variation also exists from plant
to plant.
It has a special role to play in the transport of water into plant cells. The phenomena of
plasmolysis depend upon osmosis. We have observed that water moves under the influence of
imbibition and thus water potential Ψ) must be affected by these forces. Matric potential (ΨM) is
the term commonly employed to account for all the forces causing imbibition or holding the
water in a matrix.
The potential of water in a matrix e.g. in a soil, colloid etc., can be explained as follows:
Ψ = Ψπ + Ψp + ΨM
1782― Jean Senebier demonstrated the essentiality of CO2 to produce O2 by the green plants.
1804―Nicholas Theodore de Saussure showed importance of water in photosynthesis. Light was
required for the evolution of O2.
1837―Dutrochet demonstrated the importance of chlorophyll in photosynthesis.
1842―Mayer described sun as the source of energy.
1845― Liebig indicated the involvement of CO2 in the formation of organic compounds.
1864― Sachs reported formation of carbohydrates in photosynthesis. He used half- leaf
experiment for this.
1905― Blackman proposed Law of Limiting factors.
1923― Warburg used Chlorella as a basic system for studying photosynthesis.
1924― CB van Niel showed that some bacteria use H3S instead of H2O for photosynthesis.
1939― Robert Hill performed first experiment to demonstrate that separate light reaction was
localized in chloroplasts.
1940-50― Melvin Calvin and Andrew Benson unravelled the outlines of CO2 fixation into
carbohydrates in photosynthesis. Calvin used 14CO2 in his experiments.
1941― S. Ruben and M. Kamen used 18O2 to confirm that O2 produced during photosynthesis
was derived from H2O.
1950― Ochoa and Vishniac showed that NADP+ could substitute as the hydrogen acceptor in
the Hill reaction.
1954―Kortschak described the formation of C4dicarboxylic acids.
1954―Daniel Arnon made discovery that light and of dark reactions photosynthesis were
separable.
1957― Emerson and his associates discovered Emerson effect.
1960― Woodward confirmed structure of the chlorophyll-α and synthesized it in the laboratory.
1960― Hill and Bendall showed two-step electron in transport system-photosynthesis.
1962― Fujita and Hattori investigated the effect of the various factors onformation of
phycobiliproteins in algae.
1962― Jagendorf introduced the concept of two-state phosphorylation.
1966― Hatch and Slack extended the work of Kortschak.
1967― Arnon proposed two-photosystem scheme of photosynthesis.
1968-69― Laetsch described ultrastructure of bundlesheath and mesophyllchloroplasts.
1973― Rouhani and his associates proposed pathways in CAM plant Sedum.
1980― GJ Lorimer showed that rubisco uses CO2 as a substrate and activator.
On the head drop of mucilage oozes out which helps in the capture of the insect. On the other
hand the sessile glands have basal and columellar cells bearing a distinct head of 2-8 cells. The
lateral walls of columellar cell is cutinized and thus resembles endodermis. The mature secretory
cells of all the insectivorous plants have wall ingrowths resembling transfer cells.
These are well developed on the radial walls of both the glands. The heads of the trichomes have
no well-developed cuticle and therefore secretion can easily pass through it. The glandular
trichomes show high activity of several hydrolases including esterase, acid phosphatase and
ribonuclease.
It has been shown that the chief site of activity of these enzymes was in the anticlinal or radial
walls of the head cells. It is suggested that enzymes are moved in the radial walls and then
through a poorly developed cuticle through the pores outside. If the sessile glands of Pinguicula
are stimulated secretion begins within one hour and within two to three hours it oozes out.
This secretion has a detergent characteristic and possibly wets the exoskeleton of the insect. The
glands of insectivorous plants are interesting since in them the secretion products move out of
the cuticle and the digested products move in the glands simultaneously.
In Drosophyllum also both types of glands are present. The mucilage is, however, secreted by the
outer stalked or tentacle glands. These glands are rich in dictyosomes. In fact during the period
of secretion these glands abound in dictyosomes.
The mucilage formation is respiration dependent and temperature also affects the rate of
secretion. The closure of the leaf is shown to be brought about by electrical signals which
originate in the multicellular sensory hairs. In Drosera the large tentacles on the leaf may be
regarded as large stalked glands which are not wholly epidermal in origin.
These glands have multicellular stalks and have a bundle of tracheids and a head with 2-3 cells.
The cuticle of these cells is perforated. Plasmodesmata are present between different cells in the
head. Transport of calcium has been traced in these glands. Through series of experiments it has
been demonstrated that digestive material can pass to the inner side against the direction of flow
of mucilage and digestive enzymes.
The situation in pitcher plant (Nepenthes sp.) is very interesting. The pitchers in this plant
possess different kinds of glands. These glands are similar in structure but differ in location and
function. There are ‘alluring glands’ which are multicellular and are present on the under surface
of the lid of the pitcher. These glands secrete nectar. On the inner wall of the pitcher several
digestive glands are distributed.
These glands have digestive and absorptive functions. They are more in distribution towards the
base of the pitcher. These glands also bring about a short distance transport of the material from
the pitcher cavity to the tissue. These glands also secrete several hydrolases like acid
phosphatase, esterase and ribonuclease, etc. The pitchers may have as much as one litre of
digestive fluid.
In Utricularia the leaves are modified into bladder-like traps and trap the insects in several ways.
On the outer side of the bladders there are sessile and stalked glands which secrete mucilage and
nectar in order to attract insects.
The inner wall of the bladder is lined with sessile glands which extract water from the contents
and cause tension in the bladder. Consequently water stream alone with the insects is moved in.
The door of the bladder is guarded by hair which open towards the inner side. The trapped
insects are not allowed to escape.
Cuscuta (Cuscutaceae) species are among the best known angiosperm parasites and about 180
species are reported in the genus. The parasite joins the host plant through a wedge-shaped
physiological bridge called haustorium.
In recent years development, ultrastructure and physiology of haustorium have attracted much
attention. Recent studies have shown that from the distal end of the haustorium hyphae like
structures develop which penetrate the individual cells the host cortex or pith.
These are search or contact hyphae. Of the two types, search hyphae grow inter and
intracellularly. Tripodi (1967) has studied details of the haustorium and also changes in the host
cytoplasm through electron microscope. This author reported that Golgi vesicles increased in the
host cytoplasm.
Dorr (1969) has reported several plasmodesmatal links between the protoplasts of host and
parasites. She has also reported hand like furrow at the tip of the hypha where it attaches to
host’s sieve tube membrane. Cuscuta is unique in developing a highly specialized contact with
the host phloem. Recent studies by Malik and Komal (1979) have shown that formation of
haustoria is sequential and is regulated by cellular relations within the host.
Haustoria fail to penetrate the secondary tissues, especially those which are lignified. There are
four stages in the developmental sequence of the Cuscutahaustorium and these are dependent
upon at least two conditions.
The first is the initiation of haustorialprimordium within the prehaustorium and the second is the
physical and chemical environment existing in the immediate surroundings of developing
haustorium. The haustorialprimordium has a programmed set of requirements and these must be
met with to affect hasutorium formation.
The initiation of the haustorium does not require the presence of host and that haustorial
extensions and basal xylem formation can be induced by specific chemical stimulants or even
physcialcondtions. Wolswinkel (1979) has reviewed the transport of assimilates and minerals
and the role of phloems unloading in the parasitic relationships.
Though Cuscuta is traditionally regarded as classical example of total parasite which is unable to
grow autotrophically, some reports are available in recent years which suggest the possibility of
photosynthesis by the Cuscuta vines.
In a recent review paper Malik and Singh (1980) have discussed the biochemical aspects of
parasitism by Cuscuta and discussed the available literature on the physiology of host-parasite
relationships. In the haustorium diverse types of digestive enzymes are reported. Most of these
hydrolases are not excreted enmasse into the host tissues.
Where host-parasite tissues did not contact no hydrolases were localized. Assumingly haustorial
cells contained lysosomes and the latter fused with the cell membranes and released their
contents into the host cells. Hydrolases act upon the host cells plasmalemma and destroy its
semi-permeability attributes.
As a result its turgor pressure is disturbed. When followed by the mechanical force of the
haustorium, these host cells get distrupted and are crushed due to the pressure and hence their
cytoplasm is released. The space thus made is occupied by the tip of the haustorium which
subsequently grows within the host tissues.
Apparently both enzymatic and mechanical processes were involved in facilitating the entry of
the haustorium in the host tissues. Major function of the cells in the tip of the haustorium is to
cause digestion and disruption of the host cells. Thus weakening of surrounding cells
enzymatically appears to be an essential precondition for the entry and pentration of the
haustoria.
There are several semi-parasites like Loranthus, Viscum, Arceuthobium, etc. Most of these
species grow on the tree branches and possess green leaves. Viscum species have a
dichotomously branched shoot bearing green leaves. Each branch produces a wedge shaped
haustorium which penetrates the host tissues.
There are also secondary haustoria which establish contact with the xylem of the host. It draws
water and minerals from the host tissue. Haustoria are also regarded as sucking roots. Loranthus
species are also referred to as Dandropthae sp.
They are semi-parasites growing on the branches of mango, dalbergia, fig and other species.
Here also primary haustoria penetrate the host tissue and obtain water and mineral supply
whereas leaves of the parasite are green and manufacture food material themselves.
In Punjab and United Province one of the most prominent root parasites is Orobanchecernui and
other species. This plant infests the roots of Brassica and Solanaceous members. The tall shoots
arise from the roots and bear pink flowers and white scaly leaves. Similarly species of Rafflesia
are also specialized total root parasites. Members of family Santalaceae are partial root parasites.
Kinetics of Growth:
Many investigators have measured size of an organism and have plotted the data as a function of
time. Growth of a plant or plant part characteristically passes through stages represented by an S-
shaped curve. Size versus time plot is the most direct way of handling growth data.
We can also plot the longrithum of the size as a function of time. In a size plot the units are
length, volume or weight but in the rate curve (Fig. 19-1) the units are length, volume or weight
per unit time (Fig. 19-2).
Growth curves plotted in any of the two ways have two clearly distinguishable phases (Fig. 19-
2). The first is rapid or exponential (a) phase during which the rate curve as well as size curve is
increasing. In the last phase, the size continues to increase but more slowly so that the rate
decreases (c).
The first phase has been called logarithmatic or exponential phase and the last phase is referred
to as senescence or decreasing phase. In many examples of growth curve, another phase is also
clearly evident. This is a phase of minimum growth rate or the linear phase (called by Sachs, the
grand period of growth), and it lies between the logarithmatic phase and the senescence phases.
Blackman (1919) put forward the idea that growth of a plant could be represented by the
equation
Wt = Wo.ert
where W1 is the final size (weight, height, etc. after time t), Wo is the initial size at the
beginning of the time period, r is the rate at which plant substance is laid down during time t, and
e is the base of natural logarithms. It should be noted that r is the relative growth rate as
discussed above.
Black-man pointed out that this equation also describes the manner in which money placed at
compound interest increases with time, and the term compound interest law is used to describe
such phenomena. Banks usually apply compound interest quarterly or annually so that the
increase in amount occurs at a jump. With plants or other biological systems, compound interest
is applied continuously, and size increases as a smooth curve.
From the same equation, it is also seen that the size of an organism (Wt) depends on the initial
size (Wo). It has been shown that seedling growth in seeds of different sizes follows such an
expectation, with the large seeds giving a large plant. This is true, however, only during the early
stages of growth, and if t is large, then the final size of plants from small and/or large seeds
frequently are equal.
From this equation it is seen that plant size also depends on the magnitude of r, the relative
growth rate. Blackman suggested that r might be used as a measure of the ability of a plant to
produce new plant material and called r the efficiency index. Plants with a high efficiency index
could be expected to perform better than those with a low efficiency index.
While r does differ among plant species, it is not constant during the life of aslant. Furthermore,
all parts of a plant are not equally involved in synthesizing new plant substances; some of the
material goes for storage or is catabolized. The usefulness of the efficiency index as a predictive
factor in evaluating performance or yield is limited.
Various embryological studies have shown that fruit development in several species is associated
with the pollen germination on the stigma and subsequent growth of the pollen tubes. In recent
years, it has become increasingly evident that pollination was essential for the proper growth of
the fruits.
Further, the developing seeds must be present to achieve normal fruit formation. Late Prof. J.P
Nitsch, a French plant physiologist, performed interesting microsurgical experiments in
strawberry.
In general, it is reasoned out that germinating pollen provides auxin to the pistil and in the latter
its contents progressively increase. Moreover, young developing seeds are also rich in auxin.
Nitsch demonstrated that the removal of seeds retarded the fruit development.
However, spraying of overies with auxin did overcome such a retardation. In case of
parthenocarpic fruits, seeds do not develop but abound in immature ovules.
Here the ovary develops normally even though there is no pollination and/or fertilization.
Parthenocarpy is widely known in grapes, banana, pineapple, etc. Several cucurbits and
solanaceous fruits can be made parthenocarpic by excluding pollination and spraying with auxin
or even GA3.
Figure 25-1 also shows different stages of fruit development e.g., initiation, anthesis, pollination,
fertilization, growth and maturation. These stages are accompanied by cellular and metabolic
activity. Also it is evident that early stages show cell division and the synthesis of new
protoplasm and there is high synthesis of cytokinins and gibberelins.
Metabolites are translocated from the leaves, stem, root, etc. during the developmental phase.
Beginning with zygote formation, there are two aspects of fruit development e.g. ‘pre-bloom’
and ‘post-bloom’. Fruit growth is measured in terms of volume, fresh weight, or dry weight.
Pre-bloom phase is characterized by the initiation of flower primordia while post-bloom occurs
after anthesis and consists of pollination, fertilization, fruit growth and maturation. There is
enormous amount of cell division and enlargement and cytokinins play a significant role.
However, cell division is immediately replaced by cell enlargement and IAA, GA contribute
significantly in this phase.
In order a fruit attains the edible state, it passes through different stages e.g., cell divisions, cell
enlargement, maturation and ripening. In the initial stages, there is rapid division of cells and this
is followed by enlargement of each cell.
Then the cytoplasm of the cells shifts to the periphery and vacuoles thus created are filled with
sugars. Subsequently there is increase in starch. At this stage fruit is mature and then the process
of ripening begins. The last stage is senescence when the fruit becomes susceptible to
physiological disorders and infections.
Two fruit types occur in abundance (berry and drupe) and they develop from a single ovary. In
drupes (e.g., coconut, mango, etc.) endocarp becomes hard. Sometimes several ovaries of one
flower develop into compound fruits e.g. in respberry, strawberry, etc.
In general several metabolites have been analysed in the developing fruits. In addition pigment
and flavour changes have also been traced.
In the following a brief account is given:
(i) Carbohydrates and nitrogenous materials:
In almost all the young fruits, there is abundance of chloroplasts which help in the synthesis of
sugars. These sugars are required for growth and development of fruits. In addition, much of the
needed carbohydrates come from the leaves.
A similar situation prevails with regard to nitrogen compounds. The carbohydrates accumulated
in the young fruit may act as substrate for respiration, produce organic acids or may participate
in the biosynthesis of fats, starch or even cellulose. The sour taste of many young fruits is due to
the abundance of citric acid and malic acid.
However, with the maturity of the fruit, their level decreases. Different fruits abound in different
organic acids. The accumulation of organic acids may also be accomplished by other
physiological mechanisms to be discussed elsewhere. Carbohydrates and nitrogen material is
used in protein synthesis of fruits.
As the fruit continues to grow, there is active protein synthesis accompanying it. In most of the
fleshy fruits starch content also goes up. During the ripening phase, there is a decrease in the
starch level. The type of sugar(s) prevalent in a fruit varies with the species. In mature fruits of
oranges, grapes, etc., organic acids decrease and sugars increase but this is not true of lemon.
(ii) Pigment changes:
In a maturing fruit, chloroplast is replaced by chromoplast and carotenoids. The colour of a
mature fruit depends upon the types and level of carotenoids. When exposed to light,
anthocyanins also change.
(iii) Flavouring compounds:
Basically, the flavouring compounds are aromatic esters, carboxyl compounds, alcohols, etc. The
flavour of banana, however is due to amyl acetate. The smell of citrus fruits is due to various
terpenoids, coumarins, etc. In any case, flavour and smell of fruits is an important commercial
characteristic.
The chemical composition of mature fruits varies. Fruits like apple, dates, mango, bananas have
abundant carbohydrates while those of olive, avocado store fats. In citrus fruits organic acids
accumulate. These are citric acid, malic acid (apple) or tartaric acid (grapes). In general, fruits
have low amount of proteins.