Control Systems in Plants—Chapter 39 Plants respond to environmental stimuli by: Sending signals between different parts of the plant Tracking the time of day and the time of year Sensing and responding to gravity and direction of light Adjusting their growth pattern and development I. Plant Hormones A. Research on how plants grow toward light led to the discovery of plant hormones Phototropism—growth toward or away from light Toward light is positive phototropism, away negative phototropism Results from differential growth of cells on opposite sides of a shoot Cells on the darker side elongate faster than those on the light side EXPERIMENTS LEADING TO THE DISCOVERY OF A PLANT HORMONE Charles and Francis Darwin—removed tip of the coleoptile from a grass seedling and it failed to grow toward light -tip was responsible for sensing light -since the curvature occurs some distance below the tip, the tip sends a signal to the elongating region. Peter Boysen-Jensen separated the tip from the remainder of the coleoptile by a block of gelatin, preventing cellular contact, but allowing chemical diffusion -seedlings behaved normally -impenetrable barrier was substituted, no phototropic response occurred -these experiments demonstrated that the signal was a mobile substance 1926—F.W. Went removed the coleoptile tip, placed it on an agar block, and then put the agar (without the tip) on decapitated coleoptiles kept in the dark—Fig. 39.3 -a block centered on the coleoptile caused the stem to grow straight up -if the block was placed off center, the plant curved away from the side with the block -Went concluded the agar block contained a chemical that diffused into it from the coleoptile tip, and that this chemical stimulated growth. -went called this chemical auxin B. Plant hormones help coordinate growth, development, and responses to environmental stimuli. Effects depend on site of action, stage of plant growth and hormone concentration The hormonal signal is amplified, perhaps by affecting gene expression, enzyme activity, or membrane properties Reaction to hormones depends on hormonal balance Major classes of plant hormones: 1. Auxin 2. Cytokinins 3. Gibberellins 4. Abscisic acid 5. Ethylene 1. Auxin—a hormone that promotes elongation of young developing shoots or coleoptiles a. apical meristem is a major site of auxin production b. moves from the apex down to the zone of elongation at a rate of about 10 mm per hour c. d. 1. 2. 3. 4. The acid-growth hypothesis states that cell elongation is due to stimulation of a proton pump that acidifies the cell wall—Fig. 39.5 Causes the crosslinks between the cellulose myofibrils of the cell walls to break This loosens the wall, allowing water uptake, which results in elongation of the cell Other effects of auxin affects secondary growth promotes formation of adventitious roots promotes fruit growth in many plants used as herbicides—affects dicots selectively allowing removal of broadleaf weeds 2. Cytokinins—modified forms of adenine that stimulate cytokinesis Function in several areas of plant growth: Cell division and differentiation Apical dominance Anti-aging hormones 3. Gibberellins—more than 80 natural occurring gibberellins have been identified Function in aspects of: Stem elongation Fruit growth Germination 4. Abscisic acid (ABA)—produced in the terminal bud and helps prepare plants for winter by suspending both primary and secondary growth. -Also acts as a stress hormone, closing stomata in times of water-stress thus reducing transpirational water loss 5. Ethylene—a gaseous hormone that diffuses through air spaces between plant cells -can also move in the cytosol -high auxin concentrations induce release of ethylene, which acts as a growth inhibitor Responsible for aspects of: Aging (senescence) in plants Fruit ripening Leaf abscission—loosing of leaves Analysis of mutant plants is extending the list of hormones and their functions C. Signal-transduction pathways link cellular responses to hormonal signals and environmental stimuli -Plants cells respond to hormones and environmental stimuli which are mediated by intercellular signals—signaltransduction pathway: 1. reception—detection of a hormone 2. transduction—amplification of stimulus and conversion into a chemical form that can activate the cell’s response 3. induction—the pathway step in which the amplified signal induces the cell’s specific response to the stimulus II. Plant Movements As Models for Studying Control Systems A. Tropisms orient the growth of plant organs toward or away from stimuli—result in curvatures of whole plant organs toward or away from stimuli 1. Phototropism—growth either toward or away from light—shoot tip is the site of the photoreception that triggers the growth response 2. Gravitropism—orientation of a plant in response to gravity Roots display positive gravitropism— curve down B. Shoots display negative gravitropism— curve upward 3. Thigmotropism—directional growth in response to touch Thigmomorphogenesis—developmental response to mechanical stimulation Turgor movements are relatively rapid, reversible plant responses 1. Rapid leave movements—travel from the leaf that was stimulated to adjacent leaves along the stem May be a response to reduce water loss or protect against herbivores Travel wavelike through plant at about 1 cm/sec Correlated with action potentials in animals, but thousands of times slower May be widely used as a form of internal communication since found in many algae and plants 2. Sleep movements—lowering of leaves to a vertical position in evening and raising of leaves to a horizontal position in the morning—due to daily changes in turgor pressure III. Control of Daily and Seasonal Responses A. Biological clocks control circadian rhythms in plants and other eukaryotes Circadian rhythm is a physiological cycle with a frequency of about 24 hours Probably set due to environmental signals A free running period is when an organism is sheltered from environmental cues causing the rhythm to deviate, can vary from 21 to 27 hours Most clocks cued by light-dark cycle resulting from the earth’s rotation, may take days to reset once the cues change— jet lag B. Photoperiodism synchronizes many plant responses to changes of season—a physiological response to day length Plants detect the time of year by the photoperiod 1. Photoperiodism and the control of flowering 2. Critical night length If daytime is broken by brief periods of darkness—plants will flower If nighttime is interrupted by short exposure to light—responses are disrupted and plants do not flower 3. Types of plants a. Short day plants—need long nights b. Long day plants—need only short nights c. Day neutral plants—unaffected by photoperiod IV. Phytochromes A. Phytochromes function as photoreceptors in many plant responses to light and photoperiod -phytochromes help plants measure the length of darkness in a photoperiod Phytochromes alternate between two photoreversible forms: Pr (red absorbing) and Pfr (far-red absorbing)—the interconversion between the two forms is a switching mechanism that controls various plant events 1. The ecological significance of phytochrome as a photoreceptor a. phytochrome functions as a photodetector that tells the plant if light is present b. Photoreception by phytochrome has a large effect on the whole plant even though very little pigment is present in plant cells c. Complementing phytochromes effect, other photoreceptors help coordinate B. V. a plant’s growth and development with its environment Phytochromes may help entrain the biological clock 1. Pfr gradually reverts to Pr a. occurs everyday after sunset b. Degradative enzymes destroy more Pfr than Pr c. At sunrise, the Pfr level increases due to photoconversion of Pr 2. Plants do not use the disappearance of Pfr to measure night length since: The conversion is complete within a few hours after sunset Temperature affects the conversion rate, thus, it would not be reliable 3. Night length is measured by the biological clock, not by phytochrome Plant Responses to Environmental stress A. Plants cope with environmental stress through a combination of developmental and physiological responses. B. Some plants have evolutionary adaptations that enable them to live in environments that are stressful to other plants. 1. Response to water deficit a. control systems in both the leaves and roots that help them cope with water deficits b. leaves—help the plant conserve water by reducing transpirational water loss c. roots—reduce growth 2. Response to oxygen deprivation a. waterlogged soil lacks the air spaces that provide oxygen for cellular respiration in roots b. some plants form air tubes that extend from roots to the surface, therefore O2 can reach the roots 3. Response to salt stress Excess salts in the soil may: a. lower the water potential of the soil solution causing a water deficit even though sufficient water is present b. Have toxic effect on the plant at relatively high concentrations -Many plants produce compatible solutes which are an organic compound that keeps the water potential of cells more negative than the soil solution without admitting toxic quantities of salt 4. Responses to heat stress a. Transpiration is one mechanism that helps plants respond to excessive heat and prevent the denaturing of enzymes and damage to metabolism b. Most plants will begin producing heatshock proteins when exposed to excessive temperatures—back up system to transpiration 5. Responses to cold stress a. Chilling of a plant causes a change in the fluidity of cell membranes b. Plants respond to the cold stress of chilling by altering the lipid composition of their membranes c. Subfreezing temperatures are the most severe form of cold stress because ice crystals begin to form in the plant. 6. Response to herbivores a. Plants counter excessive grazing by herbivores with both physical and chemical defense measures -physical—thorns and spines -chemical—distasteful or toxic compounds -recruit predatory animals to help defend VI. Defense against Pathogens A. Resistance to disease depends on a gene for gene recognition between plant and pathogen 1. most pathogen-plant interactions are nonvirulent 2. resistance is achieved with a precise match-up of an allele in the plant and the allele in the pathogen B. The hypersensitive response contains the infection 1. Plants infected by a virulent pathogen are capable of resisting infection through a localized chemical signaling system. The response involves the following factors: a. phytoalexins—antimicrobial compounds released from wounded cells b. activation of genes encoding PR proteins, some of which are antimicrobial and others which act as signals to adjacent cells c. Lignin synthesis and cross-linking of cell wall components, actions aimed at isolating the infection 2. If the pathogen is a virulent, the local response is more vigorous and is referred to as a hypersensitive response (HR) C. Systemic acquired resistance (SAR) helps prevent infection throughout the plant 1. HR response is local and specific, signals produced from an HR are conveyed throughout the plant. 2. These “alarm hormones” initiate a nonspecific systemic acquired resistance response to help protect uninfected tissue from a pathogen that might spread from its point of invasion