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Plant Responses to Environmental Stress
Plants have a number of mechanisms to cope
with stresses in their environment, which include such physical conditions as water (too much as well as drought), temperature (hot and cold), saline soils and oxygen deprivation, as well coping with biotic stresses such as predators and pathogens. Physical Stresses in the Environment Water Plants respond to potential dehydration by • Leaf drop, • Xeromorphic leaf structure • Leaf or stem succulence • Deep taproots • Abscisic acid monitors water condition in cells and leads to stomatal closure to minimize immediate water loss. Abscisic Acid and LEA Proteins • Synthesis of LEA proteins (late embryogenesis proteins). • LEA proteins occur naturally in maturing seeds as they desiccate for dormancy. • The LEA proteins help to stabilize the membranes and other proteins of the dehydrated cells • LEA genes can also help plants grow better during drought. Oxygen Heat • Plants have poor heat-regulating mechanisms. • Their metabolism, in particular, can be seriously impacted by hot temperatures. • Transpiration • Close stomata, and that shuts down transpiration. • Plants synthesize a class of proteins which may function to protect enzymes that would be denatured by the excess heat. These proteins are called heat-shock proteins. Cold • Plants produce more unsaturated membrane fatty acids in cold temperatures to maintain membrane fluidity needed for transport proteins. • This process works for gradual cooling. • Most plants drop fragile parts prior to the cold onset • Sudden frosts have a serious impact on plant • Ice crystals form in plant tissues when air temperatures fall below freezing • Antifreeze proteins to retard growth of ice crystals within cells. Salts • High concentrations of mineral salts decrease water
absorption.
• High concentrations of some mineral salts, such as
sodium, are also directly toxic to plants.
• Halophytes
• Most have active salt glands in leaf epidermal cells that
excrete salt. Water from the atmosphere condenses on
the salt secreted on the surface of the leaves. Mangrove Salt Excretion Salt Glands General Climate Response • Ring pattern indicates a general warming climate, with more growth (wider rings) during the past half century. • Moisture has similar effects on tree rings. • Early xylem growth Plant Responses to their Biotic Environment
• The best defense is a good offense, plants
respond to many predator attacks with a chemical offense, the secondary metabolites. • Distasteful and/or toxic to others • To prevent the toxin from killing the plant, plants store secondary metabolites in vacuoles • After predator chewing on the plant, these enter the predator’s digestive tract. • In some cases, the secondary metabolite is produced in a non-toxic form but gets converted in the predator’s digestive tract, often by bacteria, into a toxic form. • Allelopathy • Allelopathy The inhibition of growth in one species of plants by chemicals produced by another species.
• Toxins leached from leaves onto soil inhibit
germination and growth by potential competitors. • Eucalyptus secretions are so acidic that they can oxidize paint surfaces. • Many desert plants (like sagebrush) also exhibit this • Consistently the predator selects the plant with the least amount of the chemical to eat. The Predator Retaliation • Some insect predators have evolved resistance to secondary metabolites • The monarch butterfly (Heliconius) has resistance to the cyanogenic glycoside of its “host” and converts glycosides to unharmful molecules to obtain nitrogen. • Some beetles destroy the toxin-containing lactifers in leaves and feast on the tasty parts. Mechanisms for Plant Wound Responses • The secondary metabolites act as effective deterrents Growing Better A few plants respond to being eaten by herbivores by growing faster and better. … respond by producing multiple shoots adventitiously. Dandelions Response to being Overeaten – Normal Growth Pattern on Right Induced Chemical Defenses • In addition to the always-present secondary metabolites, plants have rapid responses that are induced by the action of the predator or pathogen. • These responses activate transduction pathways that lead to the production of chemical (and other) deterrents to predation and wounding. General Plant Signaling in Response to Predator/Pathogen Genetically Determined Pathogen Response • Many plants have evolved gene recognition • Plant can permit some "munching" but prohibit virulent infestations. • Both plant and non-virulent pathogens have genes that code for receptor proteins. • The plant genes "R" genes, pathogen's genes "avr" genes • The signal molecule may be produced by The pathogen, such as peptides in a bacterium Wall fragments of a fungus Cell wall components of the plant broken when the pathogen munched. • Oligosaccharins in plant cell walls may function as the recognition and signaling molecules. • Avr and R receptors are often called elicitors. • When the plant receptor protein and the pathogen receptor protein match, the plant mounts defenses against invasion by pathogen The Hypersensitive Response • The usual response mechanism with an Avr-R interaction is to produce antimicrobial agents, called phytoalexins, and PR proteins in the infected tissues. • One phytoalexin, camalexin, produced by Arabidopsis, is synthesized from the amino acid, tryptophan. • The PR proteins include enzymes that degrade the cell walls of bacterial and fungal pathogens. • One PR protein (chitinase) degrades the chitin walls of fungi. PR protein response is less rapid • H2O2 and NO are often synthesized immediately in the wound area • Cell death of the affected plant cells. • These chemicals are often toxic to the pathogen as well. • When a gene specific defense is successful, the plant typically seals off the infected area forming a necrosis, destroying both its own tissue and the pathogen. The Systemic Acquired Response • Not only can plants mount a defense in the infected area, but they also produce chemical signals in the infected area that are translocated to other parts of the plant to provide resistance to infection, as mentioned as one job of the PR proteins. This response is known as a systemic acquired response (SAR). • Salicylic acid, produced as part of the HR to wounding, functions to activate a systemic acquired response. • SAR resistance may be Shortlived, or last as long as a growing season, and Non-specific, but effective. Salicylic acid is particularly effective against some virus infections. Some plants produce the volatile methyl salicylate, which travels through air to both parts of the affected plant and to neighboring plants as a warning. • SA may also be exported to other parts of the plant initiating defense Pathogen resistant protein synthesis. Specific Immunity to RNA Viruses • Viruses infect both vertically (passed from generation to generation) and horizontally (through direct infection). • Some plants have enzymes that can form double-stranded RNA from the viral RNA, and chop the double-stranded RNA into siRNA (small interference RNA) to degrade the viral mRNA before it can be transcribed. Jasmonic Acid and Wound Response • A small peptide, systemin, is produced in the wound area in response to the predator’s saliva. • Systemin (first isolated from tomato plants) promotes fatty acids in the plasma membrane to be converted to jasmonic acid. Jasmonic acid moves through plasmodesmata to phloem sieve tubes throughout the plant and activates signal transduction pathways leading to proteinase inhibitors that bind to digestive enzymes of the predator. Dietary Defense • Some plants produce an amino acid (canavanine) that gets incorporated into larvae that die when they substitute it for an amino acid needed in protein synthesis. Using Proxies • Plants, being clever, also take advantage of other organisms to destroy and/or deter the plant predator. • In an intricate symbiosis, the Acacia tree feeds and hosts ants, which protect the tree from potential predators and competitors. If a pest (or a clothes pin)
touches the tree,
the ants swarm to
deter it or destroy it • Beans injured by predators secrete a volatile chemical that is detected by adjacent plants, which activate defense molecules.