Plant Movements

29
Rama Sisodia and Satish C Bhatla

Terrestrial plants are sessile and are incapable of changing their location. Yet, plants
can sense their environment and can respond to sensory information through visible
movements to optimize their survival, growth, and reproduction. The earliest
findings published in this field were from Charles Darwin in 1880 in his book
entitled The Power of Movement in Plants that explored the phenomenon of photot-
ropism. Plant movements can be defined as the changes in the spatial orientation or
conformation of an organ or its parts. Plant movements include movements toward
light, opening and closing of flowers, growth of developing roots in search of water
and nutrients, etc. In fact, inflorescence, fruits, and shoots are all capable of moving
by sensing changes in environment, thereby adjusting their position, function, and
behavior accordingly. Plant movements are extremely slow and often undetectable.
Classical examples of visible plant movements include the response of sensitive
plants like “touch-me-not” (Mimosa pudica) and the carnivorous Venus flytrap
(Dionaea muscipula). In addition to whole organ, the movement of single cells is
also possible which includes, for example, the movement of guard cells (which
regulates the stomatal pore size), as well as growth of the pollen tube through the
style. Such movements are caused by motor cells, driven by changes in pressure
from within the cells exerted against the cell walls. Pressure causes changes in the
dimensions of individual cells or tissues which results in distinctive movements.
Plant movements are classified as:

1. Tropic movements: These movements occur in response to environmental

signals. The direction of these movements is determined by vectorial signals,
such as gravity and light.
2. Nastic movements: These predetermined movements occur independent of the
direction of the stimulus. The location and structure of the motors driving these
movements determine the direction of these movements.

# Springer Nature Singapore Pte Ltd. 2018 907

S. C Bhatla, M. A. Lal, Plant Physiology, Development and Metabolism,
https://doi.org/10.1007/978-981-13-2023-1_29
908 29 Plant Movements

29.1 Tropic Movements

Tropic movements are initiated by external directional signals that are perceived by
the sensory receptors. These tropic responses are described as positive and negative
tropisms based on whether the movement is toward or away from the signal,
respectively. Gravitropic movements are directed by the perception of gravity by
specialized sensory cells and anisotropic response of motor cells. Phototropism is
guided by the direction of light. These include the movements of leaves and flowers
that track the position of the sun. Such movements are also called as heliotropic
movements, and the phenomenon is referred to as heliotropism. Climber plants have
tendrils that wrap around their support in response to friction when they encounter
mechanical support. Such touch (actually friction)-generated responses are called as
thigmotropic responses. External signals can also be chemical signals that direct
plant movements. Such movements are termed as chemotropic movements. They
include movement of pollen tube toward the egg cell in response to chemical signals
released from synergids. Hydrotropism refers to the growth movements in response
to unilateral stimulus of water.

29.1.1 Phototropism

Several plant movements are driven by the necessity of plants to maximize capture
of sunlight for driving photosynthesis. Light-directed movements, termed as photot-
ropism, have been a subject of investigation since Charles Darwin published his
book The Power of Movement in Plants in 1880. Among all environmental signals,
light is one of the most versatile signals and varies along four parameters, namely,
quality (wavelength), quantity (fluence, i.e., photons m
2), direction, and duration
(photoperiod). Phototropic responses can be termed as positive or negative
depending upon whether the plant responds by growing toward or away from the
light source. Aerial plant parts, including coleoptiles, hypocotyls, etc., are positively
phototropic, while the tendrils of climbing plants are negatively phototropic. Inter-
estingly leaves are plagiotropic organs orienting themselves at angles intermediate
to the direction of light. Growth of roots is mostly non-phototropic, i.e., not directed
by light. The stem of Hedera helix is negatively phototropic in the juvenile stage and
prefers shade, while the mature plant is positively phototropic. The light-driven
movements include subcellular movements as well as those of organs and whole
plant. The opening and closing of stomata in the presence or absence of light are
examples of light-driven movements at cellular level. Light is also known to direct
the growth of seedlings toward photosynthetically active radiation (PAR) as
seedlings transit from the use of stored reserve food (heterotrophic mode of nutri-
tion) to becoming autotrophic. This is exemplified by the growth pattern of etiolated
seedlings grown in dark and light (Fig. 29.1). Stem of an etiolated seedling is long
and spindle-like as it grows out in search of light. Upon exposure to light,
de-etiolation triggers reduction in shoot length and increase in radial growth.
Shade growing plants are also known to accelerate their shoot growth towards
29.1 Tropic Movements 909

Fig. 29.1 (a) An etiolated

sunflower seedling with
elongated stem and persistent
apical hook. (b) A de-etiolated
seedling with reduced stem
growth and straightening of
apical hook

ambient light. Nascent leaves are also affected by the intensity of light and reorient
themselves to maximize the interception of light. Thus, most terrestrial plants use
sunlight as signal to direct their growing organs for optimal utilization of light
source. Phototropism is a growth response to a light gradient. Under laboratory
conditions the growing plant organ bends toward unilateral light (Fig. 29.2). How-
ever, under normal light, as plant receives light from all sides, the bending is still
seen as the fluence rate varies from one side to the other, creating a gradient.
Differences in fluence rate influence phototropic response of bending of plant
organ. The light gradient across a plant varies according to the optical properties
of the tissue as well as the variation in incident light. The former properties relate to
complex interactions between light and the pigments which cause scattering, reflec-
tion, or diffraction of light as it passes through the cells. The pulvinus located at the
leaf base plays an important role in perception of the light signal for directing
movement of lamina to face the light. Relocation of chloroplasts within the meso-
phyll cells in relation to light is also mediated by light. In high light intensities, plants
prevent potential damage of the chloroplasts by aligning along the anticlinal walls,
i.e., parallel to the incident light. In low light, on the other hand, chloroplasts are
aligned along the periclinal walls, i.e., perpendicular to the incident light, in order to
maximize absorption (Fig. 29.3). The relationship between fluence and response is
best understood on analyzing fluence-response curves obtained by analyzing plant
response to varying amount (fluence) of total light (Fig. 29.4). A classical phototro-
pic response to unidirectional light shows an initial positive response (fPIPP—first
pulse-induced positive phototropism) represented by a peak called the first peak or
first positive curvature, and it occurs in response to a pulse of low fluence of light.
910 29 Plant Movements

Fig. 29.2 (a) Illumination by unilateral light causes the grass coleoptile to bend toward it; (b)
illumination provided from all sides, but higher intensity of light from right also results in bending
toward greater illumination

a b
Low Incident Light High Incident Light

Accumulation response Avoidance response

Fig. 29.3 (a) In low light conditions, chloroplasts orient along the periclinal walls perpendicular to
the incident light. (b) Under high light, the avoidance response involves chloroplast alignment
along the anticlinal walls, parallel to the incident light, thereby minimizing exposure to light

There is again a rise in the response curve that forms a second positive response
(sPIPP—second pulse-induced positive phototropism) on exposure to light of higher
fluence. The third phase of phototropic bending response (time-dependent phototro-
pism; TDP) occurs when the growing shoot is exposed to moderate to high
29.1 Tropic Movements 911

Fig. 29.4 A typical

phototropic fluence-response
plot. fPIPP first pulse-induced
positive phototropism, sPIPP
second pulse-induced positive
phototropism, TDP time-
dependent phototropism

illumination (unidirectional) over an extended period. It is important to note that the

photoreceptor responsible for transducing this light signal is located in the cytoplasm
and the chloroplast movement is mediated by cytoskeleton. Analysis of action
spectra for light mediated movement of coleoptile of plants such as Avena sativa,
Zea mays, etc. has revealed that they share a common photoreceptor—a flavoprotein
called phototropin. The absorption spectrum of phototropins shows a peak in the
blue and UV-A regions of the spectrum. Phototropins mediate a number of blue
light-induced responses such as phototropic movements, stomatal opening-closing,
and redistribution of chloroplast in response to intensity of light. Phototropins have
been studied in details in several plants including Arabidopsis thaliana, Avena
sativa, and Pisum sativum. Initially, phototropin was characterized in an Arabidopsis
mutant non-phototropic hypocotyl 1 or nph I, which is unable to respond to the
phototropic stimulus and lacks the 120 kDa membrane protein. It was later renamed
as phototropin 1 (phot1) because of its role in phototropism. Arabidopsis contains
two phototropins—phot1 and phot2—that have partially overlapping roles in
regulating phototropism. Phototropins have two segments (Fig. 29.5): the
N-terminus has the photosensory domain that contains two LOV domains that
bind with flavin mononucleotide (FMN) making the phototropin sensitive to light.
LOV domains share sequence homology to a number of eukaryotic and prokaryotic
protein motifs involved in sensing light, oxygen, and voltage (hence the name LOV).
The LOV domains consist of five antiparallel β-sheets and two α-helices that bind to
FMN. The C-terminus has a serine-threonine kinase activity. In dark phototropin is
unphosphorylated and inactive. The light signal initiates a photochemical reaction
which is perceived by phototropin. This sets up a signal transduction chain begin-
ning with the autophosphorylation of phototropin and initiates a phosphorylation
cascade. In addition to phototropin, another photoreceptor called neochrome has
been isolated from the fern Adiantum capillus-veneris that has the properties of both
912 29 Plant Movements

Fig. 29.5 PHOT1 and PHOT2 of Arabidopsis contain two LOV (light, oxygen, voltage) domains.
These bind to the cofactor flavin mononucleotide (FMN). C-terminal contains a kinase domain

Fig. 29.6 Differential auxin distribution as a response to unidirectional light leading to bending
response. Increased auxin accumulation on the shaded sides enhances growth and consequent
bending of the coleoptile toward light

phytochrome and phototropin and is regulated by both red and blue light. Both
neochromes and phototropins are known to operate in ferns, mosses, and algae.
The phototropic response involves differential growth on one side of organs that
responds to the light signal. The differential growth is mediated by lateral distribu-
tion of auxin as it moves from the apex basipetally to other areas. Upon unilateral
illumination, an unequal distribution of auxin ensues and causes auxin accumulation
and subsequent enhanced growth at the shaded side (Fig. 29.6). In Zea mays, the
immediate tip area of the shoot apex (1–2 mm) is involved in auxin production, and
29.1 Tropic Movements 913

the phototropic redistribution of the auxin occurs in the subapical region. The
asymmetric distribution of auxin by phototropin is mediated by disrupting the
distribution of auxin efflux facilitator protein called PIN. The latter is usually located
at the basal end of cells associated with xylem cells and facilitates the vertical
transport of auxin. Following phototropic stimulus, it has been experimentally
shown that the basal location of PIN is disrupted in the cortical cells on the shaded
side leading to accumulation of auxin and subsequent unequal growth and phototro-
pic bending. As much as 20% higher concentration of auxin is known in Brassica
oleracea hypocotyl on the shaded side with accompanying increase in auxin-
regulated genes including those that code for α-expansins that regulate cell
expansion.

29.1.2 Gravitropism

Gravity represents an omnipresent, constant force that acts on all organisms includ-
ing plants, directing their growth and movements (Fig. 29.7). These movements are
guided by gravity vector and are differentiated on the basis of vertical growth as
orthogravitropic, as exhibited by primary root and shoot, or by the angle with
which the organ grows (plagiotropic). The subterranean rhizomes exhibit a hori-
zontal two-dimensional movement and are termed diagravitropic. The
orthogravitropic movement in a single dimension facilitates the movement of the
primary root vertically downward in search of water and minerals. Similarly, the
young shoots grow vertically upward for optimal absorption of light and carbon
dioxide for photosynthesis, and they are termed negatively geotropic in contrast to
roots (positively geotropic). As the plant matures, the growth pattern shifts to
plagiogravitropic mode allowing the plant to expand three-dimensionally, increasing

Fig. 29.7 A variety of

gravitropic responses direct
growth and movements of
plants
914 29 Plant Movements

the supply of resources exponentially. The diagravitropic portions of the plants allow
movements in two dimensions with the outgrowth of buds and leaves in case of
shoots and adventitious roots growing out and thus expanding in the plant territory.
Gravitropic movements are also modified by light with both the presence or absence
of light and the direction of light modifying the response. Thus, young roots of Zea
mays seedlings exhibit horizontal or diagravitropic response in dark. However, in the
presence of red light, its roots grow vertically down changing their response to
positive geotropic. In other cases, red light may increase or decrease the gravitropic
response. Direction of light also modifies the gravitropic response. Unidirectional
illumination from a direction opposite to the gravity signal results in an intermediate
response curvature that depends on the magnitude and direction of the two vectors—
light and gravity. In roots, the gravitropic response is also modified by the water
potential gradient (hydrotropism) and also by mechanical stimulation (thigmotro-
pism). The ability to perceive gravity signal resides in specialized cells called
statocytes, which contain sedimentable starch grains called statoliths. Tissues that
contain statocytes are called as statenchyma. These are located in the columella cells
inside the root cap or calyptra. In stem and other aerial plant parts, the statocytes are
located in the starch sheath. Statoliths consist of a group of starch grains enclosed
within amyloplasts (Fig. 29.8). Usually amyloplasts are not mobile within the cell.
However, they can move in the direction of gravity in columella cells of the root cap
as well as in the starch sheath or endodermal cells. Amyloplasts’ mobility is evident
in the cells of the inner cortex of stems and the pulvini as well as in the motor tissue
located at the nodes of grass stem. Starch grains are heavier than the cytoplasm and
therefore tend to cluster under the influence of gravity near the lower surface (inner)
of the statocyte (force sensor hypothesis). The weight and size of starch grains
disturb the subcellular organization of the cytoskeletal elements and organelles
such as endoplasmic reticulum, creating a physiological asymmetry in the cell or

Fig. 29.8 Schematic illustration of root tip (a) and shoot (b, c) in longitudinal section showing the
direction of sedimentation of starch grains in response to gravity. (d) Statocytes containing
amyloplasts that have established a cytoplasmic asymmetry in the cell in response to gravity
29.1 Tropic Movements 915

tissue. It has been shown that position (or inclination) of statoliths (and not their
mass) is responsible for gravity perception in plant cells. Statoliths respond to
gravity like liquid, and not like a granular material. They respond to the slightest
change in angle as a liquid would do to attain a horizontal position. Thus, the
hypothesis of statocytes as “force sensors” is now replaced by “position sensor”
hypothesis. The gravity signal is perceived, transduced, and transmitted to generate a
response in the zone of elongation of growing root and shoot (Fig. 29.9). The
transmission and response of the gravitropic signal involve lateral redistribution of
auxin in the elongation zone. The horizontal orientation of the plant shoot or root
causes redistribution of auxin toward its lower side. This causes relatively more
growth toward the lower side in comparison to the other side causing the negatively
geotropic organ, such as the shoot or the coleoptile, to turn upward. In contrast, in
positively geotropic organs, such as roots, the higher concentration of auxin in roots

Fig. 29.9 Phases of

generation of gravitropic
response: perception,
transduction, transmission,
and physiological response
916 29 Plant Movements

inhibits elongation on the lower side in comparison to the upper side causing the root
to grow downward. The path of auxin transport in root is often explained with the
“auxin fountain model.” Auxin synthesized in the shoot is transported basipetally
down through the vascular tissue into the root and then acropetally toward the root
tip. In the columella, the direction of the auxin flow is reversed, and it moves
basipetally in a uniform manner into the cells of the root cortex in the zone of
elongation. However, when roots are displaced horizontally, the auxin flow in the
columella is redistributed laterally toward the lower side of the root. The higher
concentration of auxin in the lower side inhibits elongation relative to the upper side,
thus causing roots to turn down.

29.1.3 Chemotropism

External signals, including certain chemicals, are also known to drive plant
movements. The growth of the pollen tube through the style is guided by the
chemical secreted by the cells lining the style. Within the ovary the pollen tube
moves toward the egg steered by the chemical cues provided by the degenerating
synergids. These chemical signals include proteins like stigma/style cysteine-rich
adhesion (SCA)—a lipid transfer protein that is secreted by the epidermal cells lining
the style and is known to play a role in growth and adhesion of the tube in the stylar
tract. Another protein that plays a role in directing the growth of the tube is
chemocyanin (a member of the phytocyanin family of blue copper proteins).
Pollen-attracting peptides have been identified in Torenia as cysteine-rich
polypeptides, called LUREs, as attractants derived from synergid cells.

29.2 Nastic Movements

Nastic movements are nondirectional movements that are driven by growth or turgor
changes and are genetically predetermined. Autonomic internally directed nastic
movements include those shown by developing buds as they swell and open
up. Paratonic nastic motions are externally directed such as those of the leaves and
flowers in responses to changing light and temperature. Nyctinastic movements or
night movements involve folding of flowers and leaves with the onset of night. Other
nastic movements include rapid seismonastic movements and thigmonastic
movements in response to touch. Photonastic movements such as folding and
unfolding of leaves, opening and closing of flowers, etc. are synchronized by dark
to light transition. In contrast, the skotonastic movements are synchronized by the
light to dark transition. Thermonastic movements occur in response to temperature
fluctuations. For example, flowers of the plant Crocus (Iridaceae) open under
ambient temperature and close with lowering of temperature. Chemonastic
movements such as opening and closing of guard cells in response to the carbon
dioxide in the substomatal cavity are mediated by changes in the guard cell
osmoticum. Signals may also be generated by growth processes, like movement of
29.2 Nastic Movements 917

root hairs through the soil, seed germination, initiation of leaf primordia, and shoot
development.

29.2.1 Epinasty and Hyponasty

Nastic movements are also displayed by developing organs, such as leaves, petals,
and sepals, which exhibit structural and physiological differences between their
adaxial (upper) and abaxial (lower) sides. As these organs develop, they may
exhibit unequal growth expansion on the two sides, resulting in upward
(hyponastic) or downward (epinastic) growth (Fig. 29.10). Such bending
movements are mediated by unequal growth and are part of normal development
of the plant. For example, leaves of tomato plant growing in soil flooded with water
exhibit epinastic movement of leaves. These are under endogenous control, and
though the growth is irreversible, the bending of the organ is reversible. It has now
been proved that hyponastic and epinastic movements are controlled by the
differential distribution of auxin and ethylene. Similar epinastic and hyponastic
movements are also exhibited by developing vegetative and floral buds. The dome-
shaped vegetative apex initially produces leaf primordia by more rapid growth on
the lower side, which becomes convex, while the upper side becomes concave.
Such unequal growth results in the developing leaf primordia forming a sheath
around the young shoot primordium, resulting in a compact bud formation
(Fig. 29.11). As the growth proceeds, the lowermost leaves change from
hyponastic to epinastic growth. Expansion of the upper side changes the leaf
architecture from convex/concave to a typical flat leaf lamina. Elongation of the
internode results in the separation of the newly flattened leaf lamina, which now
becomes a part of the shoot. The sequential change of the leaf growth from
hyponasty to epinasty is under endogenous control and enables leaves to expand
and photosynthesize. Similarly, the transition from vegetative to reproductive
growth results in the vegetative shoot apex transforming into floral bud

Fig. 29.10 Epinastic

movement in tomato leading
to downward bending of
leaves in response to stimulus
(flooding)
918 29 Plant Movements

Fig. 29.11 Developing shoot bud with ensheathing leaf primordia that transit from epinastic to
hyponastic growth. SAM shoot apical meristem

Fig. 29.12 Geotropic movement of flower stalks during development in Papaver rhoeas L.

primordium, and instead of leaves it produces flower parts: sepals, petals, stamens,
and carpels. Since the function of flowers is to cause pollination, exposure of the
stamens and ovary, which is a prerequisite to the process, occurs with the epinastic
growth leading to unfolding of the calyx and corolla. In addition to these floral
parts, other structures also exhibit epinastic movements. These include the
epinastic movement of flower stalk of poppy plant as it curves down by differential
growth forming a hook (Fig. 29.12). As the flower opens, growth is reversed
resulting in hyponastic movement and straightening of the flower. Interestingly,
the poppy flower responds to rain by downward bending of peduncle and loss of
petal turgor which, therefore, causes closure of the flower, thereby preventing
pollen loss.
29.2 Nastic Movements 919

Fig. 29.13 (a) Nyctinastic movement of leaves in Nicotiana tabacum. Expanded/opened leaves
during daytime. (b) Clasping of upper leaves at night

29.2.2 Nyctinasty

Nyctinastic movements (Gr. nyctos, night + nastos ¼ closure) refer to the rhythmic
sleep movements driven by reversible turgor pressure changes. During the day, plant
leaves are typically in the horizontal or open position. The position changes during
the night with the leaves acquiring a closed position. Leaves of plants like tobacco,
clover, mint, etc. exhibit such movements (Fig. 29.13).

29.2.3 Thermonasty

Plant movements controlled by temperature fluctuations are called as thermonastic

movements. Flowers of Tulipa sp. (Liliaceae) and Crocus sp. (Iridaceae) open on
increasing the temperature and close on lowering the temperature by as less as
1–3  C around the ambient temperature (Fig. 29.14). Closure of perianth is mediated
by changes in the extension growth of the abaxial and adaxial surfaces of the
perianth itself in response to temperature fluctuations.

29.2.4 Thigmonasty

“Touch-me-not” (Mimosa pudica) plant with its pinnately compound leaves is a

popular example of a plant that exhibits thigmonastic movements, i.e., movements in
response to touch (Fig. 29.15a, d). The touch stimulus is perceived by the pulvinus
located at the base of each leaflet and is transmitted through sieve tubes. The
920 29 Plant Movements

Fig. 29.14 Thermonastic movement of Crocus sativus flower in response to temperature fluctua-
tion. Flower petals close upon lowering the ambient temperature by 1–3  C

stimulus includes ABA (abscisic acid), and ABA controlled ion transport leading to
their diffusion into the upper region of the pulvinus causing efflux of K+ ions and
water into the intercellular space. The resulting loss of turgidity of the cells in the
upper region of pulvinus causes the downward (closure) movement of the leaflets.
The opening of the leaves takes place by the gain of turgor with active pumping of K+
back into the cells (Fig. 29.15b, c). The inability of dark-grown plants to exhibit
seismonastic responses suggests that these movements are ATP-dependent. K+ efflux
is regulated by Ca+2 ions stored in vacuoles. An action potential is propagated from
cell to cell causing neighboring leaves to respond in a similar manner. In Arabidopsis,
four genes have been identified that are upregulated when the plant is touched
(TCH1–4). Three of these encode proteins linked to Ca+2 signaling, but TCH4 encodes
xyloglucan endotransglucosylase, which modifies the structure of the xyloglucan
components of the cell wall. It has been shown that treatment of tendrils with specific
jasmonates causes them to coil in an identical manner to the touched tendrils. This
effect was shown to be independent of ethylene and auxin.

29.3 Autonomous Movements

Interestingly there are few plant movements that are guided by both external and
internal signals. These include movements that are under the control of circadian
clock that are defined by the 24-hour diurnal rhythms of light and dark periods.
Examples of such movements include the unfolding (and refolding) of flowers
during the day, which synchronizes the opening of flowers for optimal periods of
pollination. The circadian movements are synchronized and rephrased by environ-
mental signals such as light or temperature.
29.3 Autonomous Movements 921

Fig. 29.15 (a, d) Thigmonastic movements of leaflets of “touch-me-not” (Mimosa pudica) plant.
(b, c) Mechanical stimulation activates ion channels that cause differential changes in action
potential and closure of the leaflets

29.3.1 Diurnal Movements and Circadian Rhythms

Plants are routinely exposed to diurnal cycles of light and dark and exhibit
rhythmic behavior in association with these changes. These include rhythmic
movements of leaves and petals and opening and closing of stomata that influence
the metabolic processes of photosynthesis and respiration. Flowers in the members
of Onagraceae, Cactaceae, Convolvulaceae, and Oxalidaceae exhibit spectacular
diurnal movements. Such flowers close by inward curvature of the perianth leaves
and open by reversing the curvature. These movements are mediated by growth
and take place as the flower/inflorescence develops. The rhythms alternate with a
regular periodicity of 24 h and are called as circadian rhythms (derived from
Latin, circa diem meaning approximately a day). Circadian rhythms are expressed
continuously even in the absence of external controlling factors, e.g., upon transfer
of plants from daily light and dark cycles to uniform conditions of either
922 29 Plant Movements

continuous dark or light. The rhythms are governed endogenously by an oscillator

or a biological clock that is coupled to a variety of physiological processes. The
biological clock is unaffected by temperature and can function normally under a
variety of climatic conditions. It is, therefore, known to exhibit temperature
compensation. In nature, the period of the circadian rhythms tends to be closer to
24 h (Fig. 29.16). Characteristic features define circadian rhythms. These include
the period which refers to the time between two points in the repeating cycle, e.g.,
between consecutive peak (or maxima) and trough (or minima). Phase refers to any
point on a cycle recognizable by its relationship to the rest of the cycle. Amplitude
is the degree to which the observed response varies from the mean, and the range is
the difference between the maximum and minimum values. The amplitude of a
biological rhythm can change, but the period remains unchanged. Under constant
conditions the length of the plant cycle is not influenced by any environmental
factor and is called as the free-running period. Under such conditions the rhythms
depart from the typical 24-hour period, either gaining or losing time, depending on
whether the period is shorter or longer than 24 h. However, under natural
conditions, the plants are exposed to a fluctuating environment, and, therefore,
the rhythms also exhibit the same period and are said to be entrained to its
environment. The entrainment can be brought about by a number of factors
with light being the strongest modulator. The environmental signals or cues are
called as zeitgebers, a German term for “time givers.” The environmental
zeitgebers are required to initiate the expression of the internally generated
rhythms. In case the rhythms damp out with time, an exposure to a zeitgeber is
required to restart them. It is important to note that the biological clock itself does
not damp out, rather a coupling between the endogenous oscillator and the
physiological response occurs. A single oscillator is often coupled to more than
one circadian rhythm, which are themselves often out of phase with each other. In
circadian rhythms, the operation of endogenous oscillator initiates response at a
particular time of the day. A single oscillator can be coupled to multiple circadian
rhythms, which may even be out of phase with each other. The entrainment of the
molecular clock is mediated by specific photoreceptors. Both blue and red lights
are effective in entrainment, indicating the role of both phytochromes and

Fig. 29.16 A typical circadian rhythm entrained to a 24-h light-dark cycle

29.3 Autonomous Movements 923

cryptochromes in this process. In Arabidopsis, phytochrome C is involved in clock

entrainment, whereas CRY1 and CRY2 proteins function as intermediates during
the signaling of phytochrome-mediated clock entrainment.

29.3.2 Photoperiodism

The endogenous circadian clock of the plant enables it to monitor the time of the day
and determines the specific time of the day at which a specific physiological or
molecular event has to occur. This ability of a plant to detect day length and thus
allow a seasonal response to occur is called as photoperiodism. Circadian rhythms
and photoperiodism are modulated by cycles of light and dark periods. Geographic
locations determine the length of the day and night. At equator, the length of the day
and night is equal, and as one moves toward the poles, the days become longer in
summer and shorter in winter (Fig. 29.17). Plants detect these seasonal changes and
influence their photoperiodic responses. The understanding of plant photoperiodic
responses was initiated with the work of Wightman Garner and Henry Allard in the
1920s as they conducted a series of experiments at the US Department of Agricul-
ture, Beltsville, Maryland, on a mutant variety of tobacco Maryland Mammoth. The
plants grew up to a height of about 5 m but failed to flower in the summer. However,
the plants produced flowers in winter or on artificially providing short-day
conditions during summer. It was, therefore, concluded that the length of the day
was critical in determining the flowering response and later this could also be
confirmed for a number of other plant species as well. Extensive experimentation

Fig. 29.17 The relative length of day changes from equator to poles
924 29 Plant Movements

lead to the establishment of the idea that flowering in plants is modulated by

photoperiod and plants can be classified under different categories. 1. Short-day
plants or SDPs: Plants flowering only under short days are called as qualitative
SDPs. Incase flowering is accelerated by short days these plants are called quantita-
tive SDPs. 2. Long-day plants or LDPs: Plants which flower only under long-day
conditions are called as qualitative LDPs. Incase flowering is accelerated by long
days the plants are called quantitative LDPs. Flowering in LDPs is promoted only
when the length of the day exceeds a certain duration called as critical day length.
The critical day length determines the flowering response in both LDPs and SDPs.
Long-day plants flower when the day length exceeds or the night length is less than a
critical duration in a 24-hour cycle. Short-day plants flower when the day length is
less than or the night length exceeds a critical duration in a 24-hour cycle. LDPs, for
example, many varieties of wheat (Triticum aestivum), monitor the lengthening of
the day as spring progresses toward summer, and as the critical day length is reached,
the plant flowers. SDPs, for example, Chrysanthemum morifolium, on the other
hand, flower in fall when the day length is less than a critical day length. The
absolute value of the critical day length, however, varies among plant species. A
detailed account of photoperiodism has already been provided in Chap. 25.

29.4 Mechanisms of Movement

Plant movements can be caused by a variety of mechanisms mediated by turgor

pressure changes, changing growth dynamics, movement by contraction, and change
in conformation.

29.4.1 Turgor-Mediated Movement

Plant movements are caused by a group of cells and tissues referred to as motors.
Motors may be single-celled and are responsible for movements, for example, that of
the guard cells, growth of the root hairs, and pollen tubes. Multicelled motors operate
to move entire leaves, roots, apical buds, flowers, fruits, and entire shoots. The motor
tissue is either separate from the organ that it moves or it is part of the moving and
growing tissue. For example, motor tissues driving heliotropic and sleep movements
are specialized and distinct from the organs they move. Cells of such motor tissues
extend reversibly to mediate such movements. In contrast motor tissues driving the
pollen tube growth are made up of cells that expand irreversibly. A common feature
of all motor-driven movements is that these are based on physical force exerted
against the cell walls, i.e., these movements are driven by changes in turgor pressure
within the motor cells. However, turgor-driven movements in immature cells occur
with the irreversible extension of cell walls and increase in cell volume and growth,
while in mature cells these movements result in reversible and elastic extension of
their cell walls. The variations in plant movements arise due to differences in the
29.4 Mechanisms of Movement 925

signals that initiate them as well as due to the location of the motor cells and the
mechanisms by which they respond. These movements rely on the turgor-mediated
changes in a specialized motor organ called pulvinus that occurs at the base of the
compound leaves in the families Leguminosae and Oxalidaceae. The motor cells of
pulvinus are made up of large thin-walled parenchymatous cells that use turgor-
mediated reversible changes in size and shape to alter the position of the leaf. The
opposite sides of the pulvinus are called as flexor (upper) and extensor (lower)
regions. At night, the extensor cells lose their turgor, while the flexor cells gain
turgor resulting in the petiole turning toward the stem leading to closure of the leaf.
During the day, the extensor cells regain their turgor, while the flexor cells lose their
turgor resulting in the petiole turning away from the stem axis and leaf opening
up. The relative swelling and shrinkage of the flexor and the extensor regions of the
pulvinus thus regulate the movement of the leaves or leaflets. The relative position of
the flexor and the extensor regions in the pulvinus is dependent on whether the
closure movement is acropetal or basipetal. Nyctinastic movements are defined by
blue light, phytochrome, as well as endogenous rhythms. The changes in the turgor
pressure within the pulvinus are mediated by massive redistribution of potassium ion
(K+) between the symplast and apoplast regions of both the extensor and the flexor
regions of the pulvinus. Depolarization of the membrane results in the K+ channels
to open and cause an influx of K+ ions into the extensor cells, thereby increasing their
turgor. A model integrating the role of phytochrome and secondary messengers, like
inositol triphosphate, has been proposed to explain leaf movements of nyctinastic
plants. Light signal activates the phytochrome and enhances the level of secondary
messengers, inositol 1,4,5-triphosphate (IP3) and diacylglycerol (DAG). The latter
stimulates the release of Ca+2 into the cytosol and causes phosphorylation of several
cytoplasmic proteins that further causes the extrusion of protons from the cell. A
proton motive force is generated thereof and causes the diffusion of K+ ions into
the cell.

29.4.2 Growth-Mediated Movement

Plants, being immobile, maximize their growth and survival by directing their
growing active parts toward available resources. The growing root and shoot tips
move forward through the soil and aerial environment, respectively, by means of
motor tissues located in the elongation zone just behind the apex, with newer cells
continually being added by the dividing cells of the apical meristems. Change in
direction of the actively growing plant organs can also be achieved by growth-
mediated curvature of the elongating shoot apex. Differences in growth rates in the
opposite sides of the motor tissue as well as acceleration and inhibition of cell
elongation in one region of the organ vs. the other side result in curvature and
thereby bring about change of spatial orientation of the organ. Forward movement of
root hairs, as it directs its growth around the soil particles, is affected by a shift in the
pattern of growth of root tip toward the side opposite to the one that encounters the
soil particles, thus growing around the obstacle and moving forward.
926 29 Plant Movements

29.4.3 Movement by Change in Conformation

Change in the architecture and conformation also leads to movements such as those
exhibited by the stomatal guard cells. The anisotropic structure and differentially
thickened cell walls of the guard cells cause changes in their volume, thereby leading
to opening and closing of stomata. The guard cells thus represent single-celled
motors that operate by conformational changes. Multicellular motors, such as
those that operate in the leaves of grasses, cause extensive changes in leaf confor-
mation. These include motors located on the sides of the midribs along the major
veins and which consist of a single row of large bulliform cells. As the grass leaves
dry up, these bulliform cells dry up and cause the leaf lamina to roll along its length.
When water becomes available, the cells inflate and thus flatten the leaves.

29.4.4 Movement by Contraction

In addition to movement effected by change in conformation, some movements are

caused by contraction, for example, the movement of bulky storage organs, such as
bulbs and corms, through the soil caused by specialized contractile roots. The
contractile roots are characterized by the presence of an upper unbranched
non-absorbing, thick fleshy part and a lower branched distal part. The radial expan-
sion of the fleshy part of the roots caused by the growth of the parenchyma cells of
the cortex creates a space in the soil facilitating the movement of the storage organ
(Fig. 29.18).

Fig. 29.18 Enlargement-associated movement of the Gladiolus bulb through the soil. The con-
tractile roots contract to pull down the attached bulb through the soil
29.5 Prey-Driven Movements 927

29.4.5 Twining Plants

Other interesting examples of motors are those that operate in twining plants in
which tendrils locate and attach to suitable support by forming extensive
circumnutations. The tendrils may be modified branches as in Bryonia
(Cucurbitaceae) and Passiflora (Passifloraceae) or even leaves as in Lathyrus
aphaca or Pisum sativum (Fabaceae). The leaf lamina is highly reduced in such
cases and is, in fact, limited to the midvein that functions as a tendril. The photosyn-
thetic function of the leaves is performed by stipules. The tendrils resemble the
torsion spring of mechanical watches, and differential contraction and expansion
along the two sides of the tendril establish a dorsiventral asymmetry between the two
flanks of the tendril resulting in progressive coiling of the tendril. Contraction of the
tendril below the support results in drawing the plant toward the support. The
thigmonastic response of the tendrils is rapid and highly selective. The epidermal
protuberances found in Cucurbitaceae members, such as Luffa cylindrica, represent
the receptors that elicit thigmonastic responses. The epidermal cells of the tendril are
interconnected by plasmodesmatal connections that integrate the protoplasts into a
symplast, thus allowing rapid transmission of the thigmonastic response.

29.5 Prey-Driven Movements

Several plants that lack chloroplasts are unable to follow an autotropic life and
survive as parasites on autotrophic plants or are carnivorous that derive nutrition
from their entrapped insect preys. These can be classified as:

• Holoparasites that are dependent on host plants for their organic nutrition, e.g.,
dodder (Cuscuta)
• Hemiparasites that have the ability to produce their own organic assimilates but
are dependent on the host for absorbing water and solutes from the soil, e.g.,
mistletoe (Phoradendron)

Carnivorous plants are generally found in nitrogen-deficient habitats and obtain

their nutrition by capturing and killing insects and absorbing the digested proteins of
those insects.

29.5.1 Parasitic Plants

Parasitic plants locate their prey using a variety of mechanisms. For example, plants
like the Santalaceae members—the mistletoes (Viscum album and V. cruciatum) and
Loranthus acacia and L. europaeus from Loranthaceae—produce brightly colored
and tasty fruits attracting birds. The birds eat the fruit, but the indigestible seed sticks
onto their beaks and is generally wiped off against a tree branch where it gets
deposited. On germination, the seedling invades the host tissue and uses a
928 29 Plant Movements

Fig. 29.19 Cuscuta or dodder plant coils around its host plant, drawing nutrition using a
haustorium

haustorium to absorb nutrition. The parasitic dodder plant (Cuscuta,

Convolvulaceae) is an obligate holoparasite that is dependent on its host plant to
complete its life cycle. It produces innumerable seeds that germinate, and the
seedling grows initially using the limited supply of nutrients available in the seed.
The seedling twines around a suitable host attracted by the smell of a volatile
compound or chemoattractants emitted by the host itself (Fig. 29.19). These chemi-
cal cues are volatile terpenoids such as α-pinene, β-myrcene, and β-phellandrene. As
a contact is established, the parasitic epidermal and parenchymatous cells differenti-
ate to form a secondary meristem and prehaustoria called as adhesive disk. Signals,
like mechanical pressure, osmotic potentials, and phytohormones, such as auxins
and cytokinins, initiate and control prehaustoria formation. Secretion of adhesive
substances like pectins and other polysaccharides by the prehaustoria enhances
adhesion which is further reinforced by the secretion of sticky substances such as
arabinogalactan proteins by the host plant. Host cells also respond by an increase in
the cytosolic calcium concentration initiating a signal transduction cascade. The
prehaustoria develops into parasitic haustoria that penetrate the host stem by exerting
mechanical pressure supported by degradation of host cell walls by the hydrolytic
enzymes such as pectinases and cellulases. The haustoria invade the host tissue,
establishing functional connections to the xylem and phloem of the host. Cuscuta
forms a continuous symplastic connection with its host. The haustoria form an
interface where nutrients, solutes, and carbohydrates as well as macromolecules,
such as mRNA, proteins, and viruses, are exchanged between the host and parasite.

29.5.2 Carnivorous Plants

Carnivorous plants have developed strategy to entrap their mobile prey. The preys
are attracted to the traps by means of sight or smell. Traps are made up of modified
leaves and can be passive traps or active traps driven by mechanical triggers. Plants
with mobile-active traps are the snap traps of Venus flytrap (Dionaea muscipula) and
29.5 Prey-Driven Movements 929

Fig. 29.20 Pitcher plants (Nepenthes sp.) entrap the prey in the pitcher-like traps filled with
digestive enzymes

the suction traps of bladderworts (genus Utricularia). These are triggered by the prey
and are activated by electrophysiological signaling processes. Passive traps do not
move and use slippery surfaces, sticky secretions, etc. to trap their prey. Passive traps
include those used by pitcher plants Nepenthes sp. (family Nepenthaceae), which has
a pitcher-like container, formed by leaf lamina, and is filled with digestive enzymes
secreted by specialized glands (Fig. 29.20). The prey attracted by the pitcher plant
slips into the slippery neck of the trap and with its exit blocked by the stiff epidermal
hairs pointing in one direction drowns into the enzyme soup and is digested away.
Such traps are stationery and do not involve movements. However, carnivorous
plants like Pinguicula grandiflora (family Lentibulariaceae) involve the use of
movements of the leaves to entrap the prey in a cagelike structure formed by the
folding of the leaf lamina as its cells loose turgor forming a bowl-like structure.
Similarly, the sundew plant (Drosera sp. Family Droseraceae) uses traps capable of
large-scale movements. The upper surface of the leaves in these plants bears long
flexible tentacles each of which has a mucilage and enzyme-secreting gland at the
top. The tentacles are arranged in concentric circles in the center of the lamina and
are inclined at an angle to the surface. Those that are in the center are the ones that
function to produce digestive enzymes, while those away from the center form the
trap. Mechanical stimulation generated by the trapped prey in the tentacles is
transmitted within seconds enabling rapid curvature of the tentacles toward the
center where the prey drowns in the digestive enzymes secreted by the central
tentacles forming mucilage-filled bowl. The trap of the Venus flytrap (Dionaea
930 29 Plant Movements

muscipula) is driven by turgor-mediated changes in lamina of specialized leaves that

have two identical lobes on either side of the midrib. These lobes have long stiff
bristles located perpendicular to the laminar surface (Fig.29.21a, b). Mechanical
stimulation of the bristles by the prey activates mechano-sensitive ion channels. This
causes induction of electrical signaling involving action potential, the transient
depolarization of the plasma membrane (Fig. 29.21c). The action potential mediates
the movement of the two lobes of the trap to move toward each other with the midrib
functioning as a hinge and the bristles on the opposite lobes precisely interlocking to

Fig. 29.21 (a, b) Venus flytrap (Dionaea muscipula) has leaves modified to act as traps with
bristles on edges that interlock upon excitation to entrap the prey. (c) Mechanical stimulation of
bristles causes generation of action potential that mediates the movement of the two lobes toward
each other trapping the prey
29.6 Movements for Dispersal 931

form a trap for the prey. For activation, the hairs must be touched within 20 s of each
other. Each movement generates an action potential and is recorded by the plant.
Induction of action potential causes the two lobes of the leaf to close and secrete
digestive enzymes. Aquatic plant—Utricularia (Lentibulariaceae)—is known to
possess traps with hinged trap doors. These traps are formed by leaves modified to
form flattened pear-shaped bladders attached to the submerged branches of the plant
by tiny stalks. At the entrance to the trap opening is a flap-like tissue that forms an
inward opening door. Several bristle-like hairs surround the submerged trough,
which entraps the prey preventing their exit.

29.6 Movements for Dispersal

Ability to undergo sexual reproduction and production of offsprings has been the
primary reason why angiosperms have been able to dominate earth. Plants have
devised their developmental programs to produce offsprings under optimal
conditions to ensure their survival. Also, these offsprings, comprising of either
seeds or spores, are packaged and dispersed using remarkably diverse mechanisms.
Life cycle of all terrestrial plants comprises of a sexual generation that begins with
unicellular spores produced within sporangia borne on specialized leaves called
sporophylls. The sporangia of all seed plants, i.e., angiosperms and gymnosperms,
are heterosporous, bearing numerous small male spores or microspores, also called
as pollen grains borne within microsporangia (pollen sacs), and usually a single
female spore, called megaspore located in the megasporangium. The megaspore
develops into the female gametophyte and is enclosed within the protective mega-
sporangium tissue and integument/s that forms the ovule. Upon fertilization, the
fertilized egg develops into the diploid embryo, and the ovule becomes the seed
enclosed within the ripened ovary now referred to as fruit. The fruit performs the
dual function of protecting the seed and also aids in the dispersal of seed. A
remarkable diversity of fruits is seen in angiosperms, and these are classified on
the basis of characteristics such as whether it is dehiscent or indehiscent, dry or
fleshy, etc. These features enable the fruits to facilitate seed dispersal.

29.6.1 Cohesion-Mediated Seed Propulsion

Passive dispersal of seeds is observed in plants that have dry dehiscent fruits such as
capsules of Lilium (Liliaceae) and Iris (Iridaceae). The seeds in these plants are
liberated by gravity along with the help of wind, rain, or animals. In many plants
belonging to the families Acanthaceae, Balsaminaceae, and Euphorbiaceae, ballistic
explosive movements of fruit coat at the time of dehiscence are responsible for
dispersal of seeds. Other examples include Alstroemeria (Alstroemeriaceae), Lotus
(Papilionaceae), and Bauhinia and Cytisus scoparius (Fabaceae). In Ricinus
communis the capsules dehisce explosively as the fleshy exocarp dries and the
endocarp dehisces along the three one-seeded valves with transverse fibers.
932 29 Plant Movements

Progressive dehydration causes the capsule to recurve and split open catapulting the
seeds to about 3–3.5 m away. Explosive dehiscence is also seen in the family
Acanthaceae as the two-valved capsule that is enclosed tightly within two lignified
sepals explode as they dry up. After rainfall, these sepals open up exposing the
capsule. The capsule valves are under increasing strain as they dehydrate after
maturation because of the structural differences between the outer and inner layers.
On imbibing the moisture, the valves split open explosively, and the seeds are
catapulted out of their capsules.

29.6.2 Turgor-Mediated Seed Propulsion

Plants, such as Ecballium (Cucurbitaceae), Impatiens parviflora (Balsaminaceae),

Cardamine impatiens (Brassicaceae), and Oxalis acetosella (Oxalidaceae) have
fruits that remain fleshy and turgid until they dehisce explosively. In such plants,
the changes in the turgor mediate the explosive dehiscence of the fruits to propel the
seeds out. Using this mechanism, the parasitic plant Lathraea clandestina
(Orobanchaceae) can catapult its seeds to a considerable distance of 12–20 m. The
fruits of the plant Impatiens are elongated capsules that have a fleshy pericarp with
several valves joined margin to margin. The subepidermal cells of the capsule are
large parenchymatous cells that are elongated in anticlinal orientation. During
maturation, turgor pressure builds up in these cells exerting pressure along the
fault lines (where the walls are weak) in the sutures. Eventually the strain splits
the sutures, and the separation is immediate resulting in explosive inward curvature
of the valves, thereby throwing the seeds to long distances.

Summary

• Movements in plants that occur in response to external stimuli are called as tropic
movements. Based on the nature of the stimuli, these movements can be classified
as phototropism, in response to light, and geotropism—in response to gravity.
Other tropic movements include hydrotropism and chemotropism in response to
water and specific chemicals, respectively.
• Nastic movements are independent of the direction of the stimuli and can be
internally (autonomous) or externally directed (paratonic). These include
nyctinasty or sleep movements; thermonasty, i.e., movements induced by tem-
perature variations; and seismonastic movements that require a touch stimulus to
occur.
• Central to the concept of movements in plants is the presence of cellulosic plant
wall and the positive hydrostatic pressure that builds up in the cell. Movements
can be attributed to change in growth or changes in turgor. Growth-mediated
movements are caused by changes in the length of the opposite sectors.
Relocation of auxin to a sector results in variation in growth on one side relative
29.6 Movements for Dispersal 933

to the other directing specific movement of the organ. Turgor-mediated

movements on the other hand take place by contraction of one sector and the
expansion of its opposite sector.
• In phototropism, the exposure to unilateral light affects the activity of auxin
transporters activating in preference those farthest away from the light, and the
resulting lateral transport of auxin causes the acceleration of the growth on the
shaded side.
• Several plant movements are guided on a circadian cycle monitored by the length
of the day. The universal biological oscillator functions as a “clock” and can be
easily monitored in plants grown under constant environmental conditions of
light, temperature, and humidity. Such plants exhibit autonomous movements
with a remarkable constant rhythm. Examples of circadian movements include
the sleep movements of leaves, the opening and closing of stomata, and the
diurnal opening and closing of flowers.
• Parasitic plants exhibit many types of movements for trapping prey. The active
traps of such plants have mobile elements that entrap the prey upon stimulation.
The touch-induced closure of the traps involves induction of electrical signaling
in the form of action potentials as voltage-gated ion channels cause transient
depolarization of plasma membrane. Several plants also have passive traps that
use slippery surfaces and sticky secretions to entrap their prey.
• Propulsion of seeds for dispersal is caused by passive cohesion as well as active
turgor-mediated mechanisms.

Box 29.1 Transcription Regulation of Circadian Clock in Plants

The circadian clock system of plants operates with three essential processes—
perception of the external stimulus, setting of the central clock or oscillator,
and induction of specific physiological response. The perception of the exter-
nal signal is mediated by photoreceptors such as phytochrome and
cryptochrome. These modulate the phase of the oscillator in response to the
external cues such as the day length. The final response is generated by the
plant mediated by clock-controlled genes. Several clock-controlled genes have
been investigated including CAB genes encoding chlorophyll a/b binding
proteins of the light-harvesting complex, those for small subunit of Rubisco,
etc. Transcription of CAB genes shows a cyclic increase at the beginning of the
day and peaks few hours later, and the expression later goes down as the day
proceeds. The CAB transcript level has been used to monitor circadian rhythms
at the molecular level. The use of a CAB2::luc (luciferase) reporter gene
cassette has been used to create a rhythmic light emission system facilitating
genetic analysis of the clock. Several clock-associated genes have been
isolated. These include the TOC1 (timing of CAB), CCA1 (CIRCADIAN
CLOCK ASSOCIATED 1) and LHY (LATE ELONGATED HYPOCOTYL),
CAT3 (a gene encoding enzyme catalase), and ELF3 (early flowering 3).

(continued)
934 29 Plant Movements

Box 29.1 (continued)

CCA1 is an important transcriptional regulator that binds to CAB promoter and
links phytochrome with the expression of CAB. The biological clock is based
on negative feedback system that relies on the mutual reciprocal feedback
inhibition of CCA1 and TOC1. The relative concentrations of the two rise and
fall alternately during the day. These oscillations are kept in phase by the
external light cue. The final physiological response is generated by the
switching on and off TOC1 and CCA1 regulated genes. One of the important
genes regulated by TOC1 includes the CO (Constans) gene that regulates the
flowering response.

Multiple-Choice Questions

1. Pea tendril coiling around a support is an example of:

(a) Thermotaxis
(b) Thigmonasty
(c) Thigmotaxis
(d) Thigmotropic
2. Curving of plant toward light is called:
(a) Thigmotropism
(b) Gravitropism
(c) Phototropism
(d) Thigmonasty
3. Sleep movements are mediated by turgor changes in specialized motor cells
located in:
(a) Pulvinus
(b) Stem
(c) Leaf blade
(d) Root
4. Roots perceive gravity by a mechanism that involves sedimentation of:
(a) Chloroplasts
(b) Statoliths
(c) Ribosomes
(d) Mitochondria
5. Which of the following is not true for nastic movements?
(a) Depend on the direction of the stimulus
(b) Are independent of the direction of the stimulus
(c) Genetically predetermined
(d) Driven by growth or turgor changes
Suggested Further Readings 935

6. Circadian rhythm is a biological rhythm with a cycle of:

(a) 12 hours
(b) 24 hours
(c) 36 hours
(d) 48 hours.
7. Curvature of the stem due to phototropic movement is due to uneven distribution
of:
(a) Cytokinin
(b) Auxin
(c) Phytochrome
(d) Gibberellin
8. Folding of leaflets of Mimosa pudica is associated with the efflux of:
(a) Calcium ions
(b) Magnesium ions
(c) Potassium ions
(d) Chloride ions
9. ___________ cells function as multicellular motors in the leaves of grasses
cause extensive changes in leaf conformation.
(a) Guard cells
(b) Bulliform cells
(c) Subsidiary cells
(d) Lenticels
10. Carnivorous plants entrap their prey in traps driven by:
(a) Turgor-mediated changes
(b) Growth
(c) Contraction
(d) Locomotion

Answers
1.b 2.c 3.a 4.b 5.a 6.b 7.b
8.c 9.b 10.a

Suggested Further Readings

Darwin C (1898) The power of movements in plants. D. Appleton, New York
Koller D (2011) The restless plant. Harvard University Press, Cambridge, MA/London
Trewavas A (2014) Plant behaviour and intelligence. Oxford University Press, Oxford