FISIOLOGI PERPINDAHAN TANAMAN

Tinggi tanaman, yang tetap untuk tanah tidak bisa bergerak dari tempat ke tempat. Tapi dalam tubuh
tanaman berbagai komponen protoplasma yang dalam gerakan konstan, ex., Gerakan air, mineral,
makanan dll tetapi bagian-bagian tertentu dari tubuh tumbuhan dalam menanggapi rangsangan
eksternal, menunjukkan perpindahan fisik yang disebut gerakan. tanaman uniseluler yang lebih
rendah juga menunjukkan pergerakan dari tempat ke tempat. Gerakan tersebut mungkin otonom atau
diinduksi.

The movement of plant structures in response to stimuli is very interesting. The stimuli may be in the
form of light, touch, chemicals, temperature, gravity or water. Thus the agency or factor that causes
movement is called stimulus. Not all parts of the plant act as the site of perception to stimuli. Only
certain regions or structures are capable of receiving the stimuli and such structures or organs are
called perception site. The site of perception need not be the structure that responds to movement.
In fact, in many cases the site of response and the site of perception are different. Nonetheless, the
stimulus has to cross through the plasma membrane of the cell or cells found in such structures that
receives the stimulus. Plasma membrane has all the inbuilt components and the potentiality to
receive stimulus and transmit it into intracellular mileu for proper response.

There is a time lag between the time at which the stimulus is applied and the time at which the
response begins. This time is called reaction time. It may vary and depends upon the intensity of
stimulus and the kind of response. If the stimulus is weak, there may not be any response at all, but
if the stimulus is adequate or in right quantity the response is positive. The time required to cause the
proper stimulus is called presentation time.

Once the plant body responds to the stimulus say sleeping movement (change in the position
direction), structures involves always come back to their original position. This process is called
recovery and the time required is referred to as relaxation time, which again varies from species to
species. If the stimulus is provided repeatedly the receiver structures do not respond with the same
intensity as the first instance, but it slows down. This effect is due to fatigue. Such behavior is
probably due to the loss of same material components required for the response. If the stimulation
is continued at increased frequency, the plant organs do not respond and behave as if they are dead
structures. Such a state is called Tetanus or extreme fatigue. Now it is believed that stimulation
(quantity) causes irritation at the site of perception and the product of irritation are then transported
to the site of response, where the structures respond and perform movement. All these events initiate
with signal, and the receptor that receives the signal becomes active and induces signal transduction
pathway in the cell cells. There will be a cascade of events that finally leads to the response.

Plants endowed with different structural adaptation and different potentialities exhibit different types
of movement either voluntarily or involuntarily. Based on the behavior as pattern, movements
stimulus, the plant movements have been grouped into physical movements and vital movements.

Physical Movements:

The structures involved in this type of movement are mostly dead. Xerochasy: Structures like the wall
of fruits, sporangia, capsules, have differential wall thickenings. During dry weather conditions, they
loose water to atmosphere. Because of this, the thick walls contract as a result they break open along
with the line of dehiscence, where the cell walls are thin and susceptible. Hydrochasy: Certain
structures, made up of hydrophilic substances, are capable of imbibing water as well. Due to
imbibition of water they swell. Due to imbibition of water peristomial teeth in Moss capsules, elators
in equisetum, etc. show movements. In fact such movements help in the dispersal of spores.

Vital Movements:

Plant movements due to the activity of living structures are called vital movements. They are further
classified into different kinds. In some plants, particularly unicellular algae, the entire cell moves from
place to place or from one position to the other or the protoplasm by itself shows continuous flux by
physical displacement. In others, where the plant body is fixed in the soil, certain structures show
bending or curvature movements. Furthermore, some of the movements are auto regulated and
propelled by innate mechanisms but other movements are induced by stimuli.

In this case the entire living cell is involved either in the movement of protoplasm or the entire body
of the plant cell from one place to another. These movements may be autonomous and induced.

Autonomous: Protoplasmic Streaming:

Protoplasm is not a static fluid. With all its complicated structures the entire protoplasmic fluid is in
constant sweeping motions. These streaming movements can be observed under high resolution light
microscopes. For example, in the cells of staminal hairs of tradescantia, the streaming is localized and
in each of these areas the direction is clockwise or anticlockwise. Tiny particulates are seen swept
along with the stream in a particular direction and particular directed path. Such compartmentalized
movements are known as Cyclosis. But in Elodea and other plants, the protoplasm shows uniform
movement but in one direction. One can observe chloroplasts movement along the cytoplasmic
streaming. Such movements are called Rotational Movements.

Protoplasmic streaming is due to the activity of contractile proteins found associated with other
microtubule network found within the cytoplasm. It is an active process microtubules play an
important role in such intra cellular movements. These structural elements contain motor proteins
and they perform the movements. The energy required for this process is derived from ATP molecules.
These movements help in the even distribution of chemical components. And the protoplasmic
movement across the plasmodesmata brings about the transportation of materials from one cell to
another. Auxin has been found to accelerate the rate of protoplasmic movements. Addition of
colchicine and cytochalasin B totally inhibits the movement, thereby indicating the involvement of
microtubules and microfilaments. Respiratory poisons like DPN, KCN, also inhibit the movement, thus
suggesting that these are active movements. Even amoebae show such movements. For that matter
all living cells exhibit autonomous movements.

Paratonic movements:

These movements are stimulated by external agents like light, chemical heat, etc. Hence they are
called taxis or tactic movements.

Phototactic Movement:Unicellular algae suspended in a test tube moves towards the light source to
obtain solar energy for photosynthesis. If the light is very intense, they move away from light; this
may be due to the raise in temperature. The movement in these cases is due to locomotor structures
like flagella or cilia. The beating of these structures propels the cells towards light. The flagellar
activity utilizes ATP. Hence these movements are active.

Chemotactic Movement; Certain flagellated or non-flagellated bacteria move towards the source of
food by lashing their flagella or by tumbling. In these cases, the stimulus is chemicals. Similarly, the
movement of spermatozoids in lower organisms like Bryophytes is directional, because the chemicals
released by mature archegonia provide the chemo stimulus. Sensing the chemicals, the flagellated
spermatozoids swim towards archegonia and enter the neck canal and bring about fertilization.

Thermotactic: Again lower organisms sense the temperature and move towards the compatible
temperature or move away if the temperature is incompatible. Sensing the change in the temperature
by the plants is autonomic but the cells always show directional movements.

All the above mentioned movements involve signal transduction pathway. Elucidation of these
pathways at molecular level is necessary and exciting.

Movements of Curvature:

Plants with their fixed plant body cannot move from place to place, but certain structures show
bending movements which may be directional or non-directional or they may be autonomous or
induced. However, some of the plant movements are due to the growth of the cells or due to change
in the turgidity of the cells. Based on the above features movement of curvature has been further
classified such as:

Autonomic Growth Movements:

Natatory; Plant structures like tendrils and stem tips exhibit differential growth alternately. This
results either in sideward or circular movements. Certain sub cylindrical stems (flat or angular) due to
differential growth at the sides show zig zag movement. On the other hand, cylindrical tendrils in
search of getting a hold on to a substratum, a waves of growth activity takes place around the tendril.
That is why it appears as if it is moving in circles. Such type of movements is called circumnutatory
movement. The growth activity of the above said structures is in-built and they show a rhythmic
pattern. Whether the hormonal fluxes are responsible for this type of growth movement or any other
innate mechanism involved is not known. But the treatment of these stems with ABA inhibits the
movements, indicating that these are phytohormone mediated.

Ephemeral Movements:

During the development of leaves and floral organs, the growth pattern determines the growth
direction of the structures. For example, leaves expand laterally by continuous radial divisions and
expansions. Similarly sepals and petals because of continuous growth of cells at the base on the inner
surface make the flower open. The hypocotyl hook of the bean seed straightens up because of one
sided growth due to expansion cells on that side. Such growth movements are called Ephemeral.
Once the movement reaches certain stage, the bending movement stops.

Autonomic Turgour Movement:

Desmodium gyrans (Indian telegraphic plant) and Eleiotis sorria have trifoliate compound leaves. In
the former case, central leaflet is larger and straight. But the lateral two leaflets are smaller which
show regular upward and downward movements. Such movements are observed only during
daytimes but not at night. Such rhythmic gyratory movements require about 5-8 minutes for the
completion of one cycle. This is due to autonomic turgour changes in the cells found in the swollen
pulvinus of the leaflets. How light brings about turgour changes or is it due to phytochrome mediated
responses or hormonal responses is not clear.

Paratonic Nastic Movements:

Paratonic movements are induced by external stimuli. Whatever may be the point or direction of
stimulus applied the movement of the plant structures is already predetermined and they exhibit
movement only in one direction. Most of the movements are turgour movements, but growth
movements are not uncommon. For example, the opening of hypocotyl hook of germinating bean
seedling, opening of circinately coiled leaves of ferns and cycas are the examples of growth mediated
nastic movements. Differential distribution of growth hormone in the adaxial (dorsal) surfaces of the
leaves; it is this that is mainly responsible for such growth movements. These movements are
permanent and they do not show temporary day and night fluctuations.

Photonasty:Many leguminous plants, with their pulvinous bases, show characteristic sleeping
movements and they show a rhythmic pattern of opening of leaf-lets in the day and closing in the
evening with a precision of a clock. In fact, such movements are attributed to circadian rhythm
operated by an inbuilt biological clock.

The photonastic mechanism has been explained on the basis if hormonal distribution in the cells of
the pulvinus. During daytimes, auxin is found in greater amount at the upper region of the pulvinus,
because of this the cells become more turgid and leaf lets open. The same process is reversed during
night because of the redistribution of auxin to lower side which causes folding by the way of turgour
changes.

Investigations into such movements have shown that these are phytochrome mediated, because red
and far red lights are very effective in opening and folding of leaflets. Phytochrome being an excitable
molecule in response to light it is known to bring about changes in the permeability of membranes.
Thus the turgour movements bring about so called sleeping movements. Added to this the change in
the permeability also involves the efflux and influx of K+ ions. The loading or unloading of potassium
ions causes increased or decreased DPD which acts as the driving force for the entry or the exit of
water which results is turgour movements.

Chemonasty:Drosera, Venus fly trap and such insectivorous plants have devised mechanisms to trap
insects to obtain nitrogenous compounds. These plants are found growing in boggy areas and they
are incapable of utilizing soil nitrogen.

The plant Drosera consists of leaves arranged in a rosette pattern. The leaf blades are flat and racket
shaped. The upper surface is made up of colorful glands which secrete sticky juice, where you very
high activity of golgi complex. There are a large number of sensitive tentacles spread out at the
margins. The tentacles have a sensitive broad base and a terminal glandular globosely head which
also secretes juice. In sunlight, these structures glisten like dew drops; hence the name sundew plant.
Serangga, salah jus berkilau madu, menetap pada permukaan ini di mana kaki mereka terjebak karena
jus lengket dan mereka tidak dapat melarikan diri. Sebagai serangga memiliki senyawa nitrogen dalam
tubuh mereka, mereka menyebar dan peka tentakel, karena senyawa nitrogen bertindak sebagai
sinyal, respon, mereka segera lipat pada serangga dan sama terjebak. Dalam hal ini senyawa nitrogen
yang ditemukan pada serangga memberikan stimulus. Di sisi lain, jika sepotong logam ditempatkan,
tentakel tidak melipat, bukan jika sepotong daging dijatuhkan, tentakel menutup segera. Namun,
setelah pencernaan, tentakel membuka perlahan-lahan dan proses ini memakan waktu beberapa jam.
gerakan chemonastic tersebut ditemukan di Utricularia, Venus fly trap, dll

Thermonasty: Flowers in Tulips and Crocus are very sensitive to temperature variations. Accordingly
the accessory whorls close or open. These movements are known to be due to turgour changes rather
than growth movements.

Siesmonasty: Mimosa pudica (Touch me not plant), Biophytum, Neptunia oleracea, Desmanthus are
very sensitive to touch, rather shock generated by the touch. Mimosa pudica has pinnately compound
leafless with a swollen pulvinus at the base every leaflet. Touching the leaves is believed to cause a
seismic shock to the leaves and this stimulus is transmitted all along the rachis downwards and reaches
the basal pulvinus of the leaf. The irritability caused, due to touch is actually transmitted through
sieve tubes, because these are the only structures which are capable of transmitting the stimulus as
fast as 1.5-20 cm/sec. The material basis for the transpiration stimulus has been found to be ABA and
ABA mediated ions. As the ABA ions diffuse along the rachis, it also trans-diffuses into pulvinus of the
leaflets.
The mechanism of upward movement of pinnules and downward bending of the entire leaf at the
base is now considered as due to the activity of the motor cells found in the pulvinus. Anatomical
studies indicate that at the lower region the pulvinus consist of thin walled parenchymatous cells,
which are loosely arranged with lot of intercellular spaces. Cells also contain many contractile
vacuoles. The cell membranes and vacuolar structures have contractile proteins. These vacuoles are
loaded with K+ ions when the leaflets are open. These motor cells are highly sensitive and active.
Such cellular organization is also found at the upper region of the pulvinus found at the base of the
leaflets.

When the stimulatory hormone ABA ,which is also called as stress hormone (released due to
irritability) reaches these specialized cells, they are stimulated to contract, collapse and extrude water
and K+ ions into intercellular spaces. This action brings about the collapsing of cells on this side and
the leaf downwards. Similarly mechanism is involved in the folding of leaflets upwards. The recovery
takes place within 5-10 minutes. The opening process takes place by active pumping in of K+ ions back
into cells. The K/ATPase pumps are supposed to be found in the cell membranes. This active process
requires energy. If the production of ATP is inhibited by respiratory inhibitors like DNP or KCN recovery
does not take place. If plants are kept is continuous dark, they fail to produce any siesmonastic
responses, which suggest the siesmonastic movements are ATP dependent active processes.

Paratonic Tropic Movements:

These movements are due to growth activity. The curvature movement is always directional, either
towards the stimulus or away from the stimulus. Most of these are phytohormone mediated
movements.

Phototropism: Phototropic movement is mostly exhibited by stem tips; for they always bend towards
the light source. The action spectrum of the light has been found to be at blue light. The blue light is
now known to bring about unequal distribution of active auxins than the total auxin content. The
pigment for absorption of blue light has been found to be Cryptochrome. The older view of destruction
of auxin and the lateral movements of auxins in response to light has been more or less ruled out. The
difference in the quantity of active auxin brings about differential growth and also curvature. Details
of this process have been explained elsewhere.
Photropic movement in in one of the fungi called Phycomycs blacesleeanus

The apoprotein cry in association with flavins bring about photoptropism, it gets activated by
absorbing blue light and autophosphorylation of ser/thr residues, that leads to transport of ions to
one side and brings about bending movement.

Geotropism: Growth of roots towards soil and the stem away from the soil in response to
gravitational force is an inbuilt mechanism endowed in stem tip and root tip of the plant. Curvature
movement of roots towards gravitational is termed as positive geotropism and the movement of the
stem away or against gravitational force is called negative geotropism. However in some plants,
rhizomes and runners grow parallel to the surface of the soil. Such a response is called
digeotropism. The obliquely growing stems show plagio geotropism (For details refer the Chapter
Auxin).
The differential response of stem tip and root tip to the same gravitational force is due to their
different innate structural and functional potentials. Though both structures are derived from the
same embryonic cell and possess the same genetic potential they behave differently probably this is
because of the programming of process driving the development of in such structures. The
mechanism of geotropism has been explained on the assumption that differential responses are due
to different concentrations of auxins. The concentration of auxin that promotes the growth of the
stem apex inhibits the growth of the root tip. On the other hand, the concentration of auxin that is
effective in the growth of the root tip is not adequate for the growth of the stem tip. This concept
suggests that stem apex requires higher concentration of auxins for its growth and roots require low
levels of auxins as optimal concentration for the normal growth.

If a seedling is placed on the soil horizontally, due to mass action of gravitational force, auxins move
downwards in both stem apex and root apex. In the stem apex, as more and more of auxin
accumulates at the lower surfaces the cells found in this region grow faster than the upper cells,
thus the stem curls upwards. But in roots, as more of auxin accumulates at the lower region, the
higher concentration inhibits the growth of cells found at the lower surface, but the cells at upper
surface grow faster because of lower concentration of auxin. Furthermore, the root cap being the
preceptor of gravitaropism produces ABA, which on translocation basipetally, reaches the lower cells
by sheer gravity. With the accumulation of more and more of ABA at lower cells, the growth of
these cells is further inhibited, but the cells at the upper region grow normally and thus bring about
the downward growth curvature for roots.

Chemotropism: Pollen tubes and certain fungal hyphae exhibit chemotropic movements, because
they grow towards organic nutrient rich media. In the case of pollen tubes, the direction of growth is
dictated by the chemical gradient generated by the embryo sac. This chemical gradient greatly
facilitates the growth movement of pollen tube towards the embryo sac, irrespective of the position
of ovules in the ovary.

Hydrotropism: Growth of roots towards water is called hydrotropism. Whatever may be the
positions of seedlings, as shown in the figure, the roots curl towards water. This positive growth
curvature might be due to greater water potential, probably acts as the motive force for the growth
of root tip. Stem and other structures are insensitive to water and they do not show any hydrotropic
curvature movements.

Thigmotropism: Plants with weaker stem spread around and require support for their growth.
Many of the climbers have developed hooks, pads and tendrils for getting a firm foot hold onto the
substrate; which may be a rock or a wooden stick or any such hard structure. For example, in the
case of tendrils of cucurbita and other species the terminal regions are soft, tender. The terminal
region of the tendrils consists of a number of fine pits, which are highly sensitive to touch. When the
tendril comes in contact with any supporting structures, the pits get stimulated and the same is
transmitted a few millimeters basally. This results in the differential growth of the tendril on one
side. This causes the curvature in the tendrils and finally they coil around the stem the supporting
structure. The time required for such coiling after the stimulus is just 3 to 5 minutes. Once the
initial reaction is set in at the basal region, the tendrils coil round the support and thus draw the
climber nearer to the support. It is suspected that auxins and ABA are involved in thigmotropic
curvatures. ABA inhibits the growth of the cells in the region of the contact and auxins stimulate the
growth of the cells on the opposite side and bring about eh growth curvature called coiling.