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1513058728Plantmovementsetext.Pdf Input Template for Content Writers (e-Text and Learn More) 1. Details of Module and its Structure Module Detail Subject Name <BOTANY> Paper Name <Plant Physiology II> Module Name/Title <Plant movements> Module Id <plant movements> Pre-requisites Basic knowledge about plant growth and physiology Objectives To make the students aware of the various types of growth movements seen in plants Keywords Darwin, circumnutation, phototropism, gravitropism, nastic movements Structure of Module / Syllabus of a module (Define Topic / Sub-topic of module ) <Plant movements> <Sub-topic Name1>, <Sub-topic Name2> 2. Development Team Role Name Affiliation Subject Coordinator <Dr.SujataBhargava> Savitribai Phule Pune University Paper Coordinator <Dr.SujataBhargava> Content Writer/Author (CW) <Dr. SujataBhargava> Content Reviewer (CR) <Dr. Sujata Bhargava> Language Editor (LE) <Dr. Sujata Bhargava> Management of Library and Information Network Library Science Network TABLE OF CONTENTS (for textual content) Introduction Types of movements Circumnutations Tropic movements Phototropism Gravitropism Thigmotropism Nastic movements Thigmonasty and Nyctinasty Nastic movements in carnivorous plants Summary and conclusions References PLANT MOVEMENTS Introduction Darwin in 1881 published a book called 'The Power of Movements in Plants,' in which movements are described as 'circumnutations' and the causes underlying such movements have been discussed as '.....increased growth, first on one side and then on another, is a secondary effect, and that the increased turgescence of the cells, together with the extensibility of their walls, is the primary cause of the movement of circumnutation.' Further, Darwin described how plants sense external stimuli such as light and gravity and are able to respond through directional growth- mediated movements. He mentioned that that perception of a stimulus and plant growth response do not necessarily happen in the same organ of the plant and hypothesized that a factor moves from the site of stimulus perception to the site of growth response. More than 100 years have passed since Darwin's observations and our understanding of the mechanisms underlying plant movements have also advanced. In this module we will study some of the advances made in our understanding of plant movements. Types of movements Plant movements can be classified into two main categories, namely movements that are spontaneous and those that are induced in the presence of specific stimuli. Spontaneous Management of Library and Information Network Library Science Network movements are also called nutational (or circumnutational) movements, while the induced movements, consist of tropic movements if movement occurs in the direction of the stimulus, or nastic movements, if the movement is independent of the direction of stimulus. Circumnutations These are autonomous movements mostly shown by young growing parts of a plant like hypocotyls, coleoptiles, shoots, tendrils, petioles etc. The tips of these organs outline a circular, elliptical or zigzag shape over time that may range from minutes to hours. Due to simultaneous elongation of the organ, circumnutations over a longer period are seen in the form of a helix. a b Fig 1. (a) Shoot tips of Phaseolus showing circumnutation. (b) A 3-D trace of circumnutations in shoot tips during day (lower part) and night (upper part) shows that they vary in amplitude, dependent on availability of light. (From: Stolarz, M., 2009) Circumnutation movements are ultraradian movements, with a period of occurrence that is less than a day. These movements may change in amplitude, period or direction dependent on stimuli received from hormonal applications, morphological features, temperature, light or gravity. Hence external application of gibberellic acid led to an increase in amplitude of the circumnutations in Phaseolus shoots, while the dark period increased the circumnutation in sunflower seedlings (Fig 1). The period of circumnutations also varied from several minutes minutes to hours in Phaseolus epicotyls in a temperature-dependent manner. At lower temperature of 15oC, the period of circumnutation was 27 min, while at higher temperature of 27oC, low-amplitude oscillations were Management of Library and Information Network Library Science Network observed having a period of 12 min. Hence the period is seen to become shorter with an increase in temperature, with an increase in plant age or in response to a mechanical stimulus. Two types of circumnutation periods are generally recognized; short period nutations(20–60 min long) and long period nutations(1–8 hrs). Hence an ultraradian oscillator having a broad period range may be responsible for circumnutations. The direction of circumnutation may also differ, sometimes even in the same plant. For example in Arabidopsis seedlings, the short period nutations generally occur in a clockwise direction, while the long period nutations are counterclockwise. The change in direction of the circumnutation may be induced by stimuli like gravity or touch or they may be spontaneous. Circumnutations may therefore be considered as an outcome of growth and intercellular communication, which are regulated by an oscillator (a biological clock). A model for circumnutations ascribes the movements to asymmetric plasmodesmata development, due to which the symplastic transport of growth substances, especially auxins, is irregular. Besides this, asymmetric changes in turgor due to alterations in ion content may also be responsible for these movements. Changes in intracellular K+ and Ca2+ ion concentrations were shown to change the amplitude and period of circumnutation probably by altering the electric potential across the plasma membrane. The function of circumnutation is obvious in climbing plants, which seek mechanical support by nutations of the young shoots and tendrils. Circumnutations are also thought to stabilize hypocotyls during elongation and inhibition of circumnutations is thought to be the first response of plants subjected to stress (before growth ceases). Tropic movements Tropic movements are not spontaneous like circumnutations, but are directional growth mediated movements in response to external stimuli like light, gravity, touch etc. They enable plants to perform better, for example by directing growth of shoots towards light, or directing root growth towards water. 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. Management of Library and Information Network Library Science Network Phototropism It is the directional curvature of organs in response to differential light and has been mainly observed in young shoots of seedlings. This response ensures that maximum light is available to the plant for photosynthesis. Roots are negatively phototropic, which means that they grow away from light and possibly meant to ensure that the roots grow into soil for water and nutrient absorption. The direction of phototropic curvature depends on the direction of light stimulus and occurs due to differential cell elongation rates, such that the side receiving less light show more elongation than the side facing the light source, hence causing bending of the stem towards light. This differential elongation was shown to arise due to differential accumulation of the plant hormone auxin on the two sides of the stem (Went and Thimann, 1937). This differential auxin accumulation is due to lateral movement of auxin from the illuminated side to the shaded side. PIN proteins, which are components of the auxin efflux carrier, play an important role in this lateral auxin transport. Fig 2. Lateral movement of auxin due to directional exposure to blue light. (a) Differential auxin accumulation across stem during phototropic curvature (b) Lateral auxin transport in response to light exposure as in (a). Hence directional light has to be perceived by the stem and the signal transduced to bring about lateral auxin transport. Phototropins are blue light photoreceptors and are coded for by two genes PHOT1 and PHOT2. The phototropins consist of two photosensory domains that show similarity to proteins responsive to light, oxygen or voltage (LOV domains) and a phosphorylating Management of Library and Information Network Library Science Network kinase domain. The chromophore is Flavin mononucleotide (FMN), which is bound non-covalently to the two LOV domains. In the presence of blue light of 447nm, FMN excitation brings about a covalent bond formation between FMN and the SH group of a cysteine present in the LOV2 domain to form the LOV 390 form. This brings about an activation of the kinase domain, which autophosphorylates the protein. LOV390 is the active form of phototropin which is reversed to LOV447 in dark. Fig: 3(a) Phototropins consist of two photosensory (LOV) domains with FMN as the chromophore. (b) Activation of phototropin by blue light causes bond formation between FMN and the LOV protein, which activates the kinase leading to autophosphorylation. (From: Christie, 2007). Phototropins are located in the plasma membrane in dark, but are rapidly internalized in the presence of blue light, indicating that the phosphorylated form of phototropins dissociates from the membrane. This leads to a reduction in blue light sensing ability of cells. Phototropins are thought to play a role in the formation of differential auxin gradients. Though the exact mechanism of
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