An Overview of Plant Defenses against Pathogens and Herbivores

Plants represent a rich source of nutrients for many organisms including bacteria, fungi, protists, insects, and vertebrates. Although lacking an immune system comparable to animals, plants have developed a stunning array of structural, chemical, and protein-based defenses designed to detect invading organisms and stop them before they are able to cause extensive damage. Humans depend almost exclusively on plants for food, and plants provide many important non-food products including wood, dyes, textiles, medicines, cosmetics, soaps, rubber, plastics, inks, and industrial chemicals. Understanding how plants defend themselves from pathogens and herbivores is essential in order to protect our food supply and develop highly disease-resistant plant species. This article introduces the concept of plant disease and provides an overview of some defense mechanisms common among higher plants. A close examination of plant anatomy is presented, as well as some of the ecological relationships that contribute to plant defense and disease resistance. Special care has been taken to illustrate how products used in everyday life are derived from substances produced by plants during defense responses.

Plant Disease and Resistance
Broadly defined, disease is any physiological abnormality or significant disruption in the “normal” health of a plant. Disease can be caused by living (biotic) agents, including fungi and bacteria, or by environmental (abiotic) factors such as nutrient deficiency, drought, lack of oxygen, excessive temperature, ultraviolet radiation, or pollution. In order to protect themselves from damage, plants have developed a wide variety of constitutive and inducible defenses. Constitutive (continuous) defenses include many preformed barriers such as cell walls, waxy epidermal cuticles, and bark. These substances not only protect the plant from invasion, they also give the plant strength and rigidity. In addition to preformed barriers, virtually all living plant cells have the ability to detect invading pathogens and respond with inducible defenses including the production of toxic chemicals, pathogen-degrading enzymes, and deliberate cell suicide. Plants often wait until pathogens are detected before producing toxic chemicals or defense-related proteins because of the high energy costs and nutrient requirements associated with their production and maintenance.
Plant Pathogens: Sneak Attack versus Brute Force

Many plant pathogens act like “silent thieves” who want to steal money locked inside of a bank vault. These thieves use specialized tools designed to disable the bank’s security system and unlock the vault without being detected. In a similar way, many pathogens establish intimate connections with their hosts in order to suppress plant defenses and promote the release of nutrients. Pathogens that keep their host alive and feed on living plant tissue are called biotrophs. Examples of biotrophic pathogens include the powdery mildew fungus Blumeria graminis and the bacterial rice pathogen Xanthomonas oryzae. Other pathogens resort to brute force like thieves who blast open a bank vault with explosives. These pathogens often produce toxins or tissuedegrading enzymes that overwhelm plant defenses and promote the quick release of nutrients. These pathogens are called necrotrophs, and examples include the gray mold fungus Botrytis cinerea and the bacterial soft-rot pathogen Erwinia carotovora. Some pathogens are biotrophic during the early stages of infection but become necrotrophic during the latter stages of disease. These pathogens are called hemibiotrophs and include the fungus Magnaporthe grisea, the causative agent of rice blast disease.

Figure 1. Powdery mildew on a maple leaf

Figure 2. Rice blast disease on rice Figure 3. Blackleg soft-rot on leaves a potato tuber

resistance. Basal resistance can be triggered when plant cells recognize Figure 5. Although drastic compared to basal resistance. for example) are capable of recognizing the bacteria and limiting disease via resistance (an incompatible response). The HR is typically more pathogen-specific than basal resistance and is often triggered when gene products in the plant cell recognize the presence of specific disease-causing effector molecules introduced into the host by the pathogen. plants may respond with another line of defense: the hypersensitive response (HR). lipopolysaccharides. A plant species that does not show disease when infected with a pathogen is referred to as a non-host plant species for that pathogen. two outcomes are possible: A compatible response is an interaction that results in disease. Disease resistance exists as a continuum of responses ranging from immunity (the complete lack of any disease symptoms) to highly resistant (some disease symptoms) to highly susceptible (significant disease symptoms). Pathogens have developed countermeasures that are able to suppress basal resistance in certain plant species. For example. . When a pathogen is capable of causing disease on a particular host species. is the first line of pre-formed and inducible defenses that protect plants against entire groups of pathogens. If a pathogen is capable of suppressing basal defense. syringae. Non-pathogens as well as pathogens are capable of triggering basal resistance in plants due to the widespread presence of these molecular components in their cells. The HR is characterized by deliberate plant cell suicide at the site of infection. These surveillance systems are linked to specific pre-programmed defense responses. some tomato cultivars show disease when infected with the bacterial pathogen Pseudomonas syringae (a compatible response). fungi. Although a particular plant species may be a susceptible host for a particular pathogen. viruses. The result is that living plant cells become fortified against attack. Figure 6. Bacterial flagella are often microbe-associated molecular patterns (MAMPs) including specific recognized by plants during basal proteins. HR lesion on an Arabidopsis leaf. but others (cultivar Rio Grande. and the leaf on the right is resistant. such as the saprophytic bacterial species Pseudomonas putida.Most biotrophic and hemibiotrophic pathogens can only cause disease on a relatively small group of host plants because of the slightly different set of specialized genes and molecular mechanisms required for each host-pathogen interaction. Tomato leaves exposed to the bacterial pathogen P. For example. while an incompatible response is an interaction that results in little or no disease at all. Basal resistance. and microscopic worms called nematodes are capable of inducing the HR in plants. also called innate immunity. the HR may limit pathogen access to water and nutrients by sacrificing a few cells in order to save the rest of the plant. are referred to as non-pathogens. Organisms that do not cause disease on any plant species. brome mosaic virus (BMV) infects grasses such as barley but not legumes. Surveillance and Detection of Microbial Pathogens Figure 4. and cell wall components commonly found in microbes. Bacteria. The host range refers to the plant species on which a pathogen is capable of causing disease. The leaf on the left is diseased. Plants have developed multiple layers of sophisticated surveillance mechanisms that recognize potentially dangerous pathogens and rapidly respond before those organisms have a chance to cause serious damage. some individuals may harbor genes that help recognize the presence of the pathogen and activate defenses.

All plant cells have a primary cell wall. Plants can recognize these foreign molecules and respond by digesting the genetic strands into useless fragments and halting the infection. These chemicals may repel harmful insects or attract beneficial predators that prey on the destructive pests. compared to the thick secondary cell walls of oak which give strength and flexibility to the wall. Feeding on one part of the plant can induce systemic production of these chemicals in undamaged plant tissues. It provides an excellent structural barrier that also incorporates a wide variety of chemical defenses that can be rapidly activated when the cell detects the presence of potential pathogens. The cell wall is a major line of defense against fungal and bacterial pathogens. a process analogous to the memory of vertebrate immune systems. and once released. invasion and infection. but disease symptoms may eventually disappear if RNA silencing is successful. cell walls of red pepper cells (A) are relatively thin These chains are bundled into fibers called microfibrils. these chemicals can act as signals to neighboring plants to begin producing similar compounds. plant tissues may become highly resistant to a broad range of pathogens for an extended period of time. Detection of Insect Herbivores Mechanical damage caused by insects is not generally considered “true” plant disease although plants have developed surveillance systems designed to recognize insect pests and respond with specific defense mechanisms. The primary cell wall consists mostly of cellulose. In response. Many viruses produce double-stranded RNA or DNA during replication in a host cell. a process called recovery. the plant may retain a template of the digested genetic strand that can be used to quickly respond to future attack by similar viruses. Comparison of cell wall types. and homoterpenoids. plants can defend themselves against viruses by a variety of mechanisms including a sophisticated genetic defense system called RNA silencing. which provides structural support and is essential for turgor pressure. These substances are gaining favor in the agricultural community because they are much less toxic to humans and wildlife than fungicides or antibiotics. a complex Figure 7. This phenomenon is called systemic acquired resistance (SAR) and represents a heightened state of readiness in which plant resources are mobilized in case of further attack. Plants that are infected with viruses will often exhibit chlorosis and mottling. and their protective effects can last much longer. Production of these chemicals exacts a high metabolic cost on the host plant. Structural Defenses The Plant Cell All plant tissues contain pre-formed structural barriers that help limit pathogen attachment. including monoterpenoids. sesquiterpenoids. In addition. Researchers have learned to artificially trigger SAR by spraying plants with chemicals called plant activators. and cotton plants produce volatiles that attract predatory wasps when damaged by moth larvae. Lima beans and apple trees emit chemicals that attract predatory mites when damaged by spider mites. The walls of pear fruit stone cells (C) are so wall may also contain two groups of branched thick that the cell lumen is barely visible. Plants can distinguish between general wounding and insect feeding by the presence of elicitors contained in the saliva of chewing insects.Once the hypersensitive response has been triggered. and many also form a secondary cell wall that develops inside of the primary cell wall after the cell stops growing. In addition to the hypersensitive response. For example. The primary polysaccharide consisting of thousands of glucose monomers linked together to form long polymer chains. The cell wood (B). plants may release volatile organic compounds (VOCs). so many of these compounds are not produced in large quantities until after insects have begun to feed. wheat seedlings infested with aphids may produce VOCs that repel other aphids. polysaccharides: cross-linking glycans and pectins. Pectins form hydrated gels that help “cement” neighboring cells together and regulate the water content . Cross-linking glycans include hemicellulose fibers that give the wall strength via cross-linkages with cellulose.

Soft-rot pathogens often target pectins for digestion using specialized enzymes that cause cells to break apart: these organisms are extremely common. are polysaccharide polymers that impede cellular penetration at the site of infection. Some plant cells are highly specialized for plant defense. There are many classes of idioblasts including pigmented cells. suberin. and they serve as a signal to neighboring cells that an attack is underway. Plant cells also respond to microbial attack by rapidly synthesizing and depositing callose between the cell wall and cell membrane adjacent to the invading pathogen. and anyone who has seen fruits or vegetables become brown and “mushy” have seen these pathogens in action. However. Grasses and sedges contain rows of silica cells in their epidermal layers which give strength and rigidity to the growing leaf blades and deter feeding by chewing insects. Lignin is the primary component of wood. Many cell walls also contain lignin. fruits. It is the first line of defense against invading pathogens and consists of both specialized and unspecialized cells. When a plant cell detects the presence of a potential pathogen. Reactive oxygen molecules also help strengthen the cell wall by catalyzing cross-linkages between cell wall polymers. an important defense against many fungal pathogens that require standing water on the leaf surface for spore germination. Some stinging cells contain prostaglandins. and roots of plants until they undergo considerable secondary growth. Sclereids are irregularly-shaped cells with thick secondary walls that are difficult to chew: the rough texture of pear fruit (Pyrus spp. floral parts. seeds.of the wall. Crystalliferous cells contain crystals of calcium oxalate that may tear herbivore mouthparts when chewed and can be toxic if ingested. Callose deposits. and these are often produced as part of the induced basal defense response. and waxes are fatty substances that may be deposited in either primary or secondary cell walls (or both) and outer protective tissues of the plant body. Pigmented cells often contain bitter-tasting tannins that make plant parts undesirable as a food source. Figure 8. The epidermal cells of aerial plant parts are often covered in a waxy cuticle that not only prevents water loss from the plant. a heterogeneous polymer composed of phenolic compounds that gives the cell rigidity. Humans and pets who chew the leaves of these plants may experience a burning sensation in the mouth and throat that is often accompanied by swelling. biting taste. stems. For these reasons. sclereids. Stinging nettles (Urtica dioica) produce stinging cells shaped like hypodermic needles that break off when disturbed and inject highly irritating toxins into herbivore tissues. waterresistant cuticle that helps prevent water loss from the plant. Idioblasts (“crazy cells”) help protect plants against herbivory because they contain toxic chemicals or sharp crystals that tear the mouthparts of insects and mammals as they feed. hormones that amplify pain receptors in vertebrate animals and increase the sensation of pain. some fungal pathogens including Fusarium solani produce cutinases that degrade the cuticle and allow the fungi to penetrate the epidermis. Cell walls contain proteins and enzymes that actively work to reshape the wall during cell growth yet thicken and strengthen the wall during induced defense. and an inability to speak. Cutin.) is caused by thousands of sclereid stone cells that can abrasively wear down the teeth of feeding animals. The hydrophobic nature of the cuticle also prevents water from collecting on the leaf surface. The epidermis is a protective layer of cells that is usually covered with a thin. Young red wines often contain high levels of tannins that give wine a sharp. Members of the genera Philodendron and Dieffenbachia are very common tropical house plants that contain large amounts of these cells. including bark. Plant Tissues and Specialized Appendages The epidermis constitutes the outermost protective tissue system of leaves. . The cuticle can be relatively thin (aquatic plants) or extremely thick (cacti). and cell walls that become “lignified” are highly impermeable to pathogens and difficult for small insects to chew. crystalliferous cells. species of Dieffenbachia are commonly called dumb cane. choking. and silica cells. but also prevents microbial pathogens from coming into direct contact with epidermal cells and thereby limits infection. called papillae. enzymes catalyze an oxidative burst that produces highly reactive oxygen molecules capable of damaging the cells of invading organisms.

Outer bark (phellem) is an excellent example of a preformed structural barrier that contains high Figure 10. Stomatal pore size is highly regulated by plants. The simplest terpenoid is the hydrocarbon isoprene (C5H8). These compounds usually belong to one of three large chemical classes: terpenoids. the “thorns” on the stem of rose plants (Rosa spp. insects from reaching the living cells underneath. Primary metabolites are substances produced by all plant cells that are directly involved in growth. diterpenoids (four units). and nucleic acids. Figure 11. The velvety appearance of dusty miller (Senecio cineraria) is caused by thousands of tiny trichomes covering the plant’s surface.) are neither true thorns nor spines: they are actually outgrowths of the epidermis called prickles. sesquiterpenoids (three units). Trichomes (“leaf hairs”) are specialized epidermal cells found on aerial plant parts that may provide both physical and chemical protection against insect pests. amino acids. or reproduction. a volatile gas emitted during photosynthesis in large quantities by leaves that may protect cell membranes from damage caused by high temperature or light. monoterpenoids consist of two isoprene units. Chemical Defenses Plant chemicals can be divided into two major categories: primary metabolites and secondary metabolites. development. Many cacti produce thorn-like structures that are actually modified leaves or parts of leaves (e.g.000 compounds described.Interspersed among the many unspecialized cells of the epidermis are guard cells which regulate gas exchange through small openings called stomata. and triterpenoids (six units). For example. Thorns are modified branches that protect plants from grazing vertebrates. Terpenoids are classified by the number of isoprene units used to construct them. and include the honey locust tree (Gleditsia triacanthos). Trichomes on the surface of soybeans (Glycine max) prevent insect eggs from reaching the epidermis and the larvae starve after hatching. Terpenoids (terpenes) occur in all plants and represent the largest class of secondary metabolites with over 22. such as in the barrel cactus (Ferocactus spp. Note the small rodshaped bacteria. phenolics. and guard cells can Figure 9. proteins. and glandular trichomes in potato and tomato secrete oils that repel aphids. In woody plants. tomato leaf. Isoprene . the periderm replaces the epidermis on stems and roots. Stomate on the surface of a participate in defense by closing in response to the presence of MAMPs. Examples include sugars. stipules) called spines which serve similar purposes. The hook-shape of snap bean (Phaseolis vulgaris) trichomes impale caterpillars as they move across the leaf surface. Phaseolis vulgaris bean leaf surface with stomata and amounts of water-resistant suberin and prevents many pathogens and uniseriate trichomes. and alkaloids. These pores allow carbon dioxide to enter the leaf for use in photosynthesis while restricting excessive water loss from the plant.).. Botanically speaking. Secondary metabolites are not directly involved in growth or reproduction but they are often involved with plant defense.

). condiments. and perfumes are made using essential oils that function as insect toxins in plants but are relatively harmless to humans. oregano (Origanum spp. Citronella is an essential oil isolated from lemon grass (Cymbopogon citratus). Digitalis caterpillars become adult butterflies.). including the insecticides permethrin and cypermethrin. Diterpenoids include gossypol. thyme (Thymus spp. Many spices. The caterpillars store these toxins safely within their bodies. they are highly poisonous to most predatory birds purpurea that eat them.and beta-pinene. including humans.). Azadirachtin Figure 15. and can cause heart attacks if ingested in high quantities.). Mint plants (Mentha spp. Some herbivores have overcome the dangerous effects of cardiac glycosides and actually use these toxins for their own benefit. Gossypol Figure 14. Black walnuts (Juglans nigra) produce juglone.). . Pine tree resin contains large quantities of the monoterpenoids alpha. called pyrethroids. Azadirachtin is a very powerful limonoid isolated from neem trees (Azadirachta indica): some insects are repelled by concentrations as low as a few parts per million. Triterpenoids are similar in molecular structure to plant and animal sterols and steroid hormones. and when the Figure 16. Figure 13.) produce large quantities of the monoterpenoids menthol and menthone which are produced and stored in glandular trichomes on the epidermis. Pyrethrins are monoterpenoid esters produced by chrysanthemum plants that act as insect neurotoxins. When produced by plants such as spinach (Spinacia oleracea). these compounds give the organic solvent turpentine its characteristic sharp odor. a chemical that interferes with the normal development of other plants. savory (Satureja spp.g.). which are used medicinally in small quantities to treat heart disease in people. tomato). Phytoectysones are mimics of insect molting hormones. which are potent insect repellents. Digoxin Insects are not the only herbivores that feed on plants. especially members of the nightshade family (e. Essential oils often function as insect toxins and many protect against fungal or bacterial attack. they disrupt larval development and increase insect mortality.). Foxglove (Digitalis purpurea) is the principal source of the cardiac glycosides digitoxin and digoxin.) which contains high amounts of these toxins in the milky latex of their sap. Figure 12. Monoterpenoids are not just used as insecticides. seasonings.). a terpenoid produced by cotton (Gossypium hirsutum) that has strong antifungal and antibacterial properties. rosemary (Rosmarinus spp..Monoterpenoids and sesquiterpenoids are the primary components of essential oils. Examples include peppermint and spearmint (Mentha spp. The fresh scent of lemon and orange peels is the result of a class of triterpenoids called limonoids. it contains high limonoid levels and has become a popular insect repellent in the United States due to its low toxicity in humans and biodegradable properties.). black pepper (Piper spp. Triterpenoids such as cardiac glycosides are highly toxic to vertebrate herbivores. Monarch butterfly caterpillars feed almost exclusively upon milkweed (Asclepias spp. basil (Ocimum spp. cinnamon (Cinnamomum spp. sage (Salvia spp. Many commercially available insecticides are actually synthetic analogues of pyrethrins. which are highly volatile compounds that contribute to the fragrance (essence) of plants that produce them.). and bay leaf (Laurus spp.

Tannins are toxic to insects because they bind to salivary proteins and digestive enzymes including trypsin and chymotrypsin resulting in protein inactivation. The wheat pathogen Gaeumannomyces graminis is unable to infect oats that contain avenacins. They are produced primarily via the shikimic acid and malonic acid pathways in plants. Phytoalexins are isoflavonoids with antibiotic and antifungal properties that are produced in response to pathogen attack. rigid. Fusarium oxysporum. Psoralen. which greatly increase the absorption of certain drugs into the bloodstream from the intestines. These toxic molecules disrupt pathogen metabolism or cellular structure but are often pathogen specific in their toxicity. Because it is insoluble. and include a wide variety of defense-related compounds including flavonoids. Phenol. phytoalexins. although primary walls can also become lignified. Cyanin glycoside. Lignin is a highly branched heterogeneous polymer found principally in the secondary cell walls of plants. which contributes to rapid cell death. Phenolics Phenolics are another large class of secondary metabolites produced by plants to defend themselves against pathogens. fruits. and the leaves of deciduous plants in fall. It consists of hundreds or thousands of phenolic monomers and is a primary component of wood. rishitin produced by both tomatoes and potatoes (the Solanaceae family). lignin provides an excellent physical barrier against pathogen attack. The sharp taste of red wine is caused by grape tannins binding to salivary proteins in the mouth which results in protein coagulation. produced by Arabidopsis thaliana. tannins. anthocyanins. Figure 17. They are activated by ultraviolet light and can be highly toxic to certain vertebrate and invertebrate herbivores due to their integration into DNA. Furanocoumarins are phenolic compounds produced by a wide variety of plants in response to pathogen or herbivore attack. Some Figure 20. In fact. and furanocoumarins. Medicarpin. and virtually indigestible. These substances have detergent (soap-like) properties and disrupt the cell membranes of invading fungal pathogens. grapefruit juice contains small quantities of furanocoumarins. a class of triterpenoid saponins. Insect herbivores that ingest high amounts of tannins fail to gain weight and may eventually die. and camalexin. an Figure 19.Saponins are glycosylated triterpenoids (triterpenoids with attached sugar groups) that are present in the cell membranes of many plant species. Flavonoids are one of the largest classes of phenolics. the simplest phenolic compound Figure 18. Anthocyanins are responsible for the showy colors of many plants and are present in high concentrations in flowers. some fungal pathogens have developed counter-measures to these plant defenses: Botrytis cinerea. lignin. Examples include medicarpin produced by alfalfa (Medicago sativa). and Septoria lycopersici are all capable of degrading saponins and causing disease in susceptible saponin-producing plants. However. a anthocyanin phytoalexin Tannins are water-soluble flavonoid polymers produced by plants and stored in vacuoles. a medicines carry warning labels cautioning people to avoid drinking grapefruit juice furanocoumarin while taking the drugs in order to avoid an accidental overdose. . Anthocyanins are colorful water-soluble flavonoids pigments produced by plants to protect foliage from the damaging effects of ultraviolet radiation.

and cocoa (Theobroma cacao). Proteins and Enzymes Many plants and seeds contain proteins that specifically inhibit pathogen and pest enzymes by forming complexes that block active sites or alter enzyme conformations. and proteinase inhibitors. Theobromine Members of the nightshade family (Solanaceae) produce many important alkaloid compounds. They include defensins. proteins require a great deal of plant resources and energy to produce. Glucosinolate Cyanogenic glycosides are a particularly toxic class of nitrogenous compounds that break down to produce hydrogen cyanide (HCN). but they are stored in separate compartments or tissues within the plant. Glucosinolates. Allelopathy allows one plant species to “defend” itself against other plants that may compete for growing space and nutrient resources. lysine. it has been used medicinally by humans in small amounts as a pupil dilator and antidote for some nerve gas poisonings. tyrosine. Figure 21. It is toxic to both insects and fungi. the enzymes and substrates mix and produce lethal hydrogen cyanide. Capsaicin Figure 25. Caffeine Figure 23. are sulfur-containing compounds synthesized by members of the mustard family (Brassicaceae) and produce cyanide gas when broken down by enzymes called thioglucosidases. It is released when herbivores graze on the leaves and break open the vacuoles. and nicotine. and many of these substances have powerful effects on animal physiology. including glycosidases and hydroxynitrile lyases. and tryptophan. Although it is toxic in large quantities. lectins. phenolics. They are derived from the amino acids aspartate. cocaine. amylase inhibitors. Atropine Figure 26. and alkaloids.Nitrogen Compounds Alkaloids are a large class of bitter-tasting nitrogenous compounds that are found in many vascular plants and include caffeine. Atropine is a neurotoxin and cardiac stimulant produced by the deadly nightshade plant (Atropa belladonna). Capsaicin and related capsaicinoids produced by members of the genus Capsicum are the active components of chili peppers and produce their characteristic burning sensation in hot. In fact. consequently. high levels of caffeine produced by coffee seedlings can even inhibit the germination of other seeds in the vicinity of the growing plants. a lethal chemical that halts cellular respiration in aerobic organisms. a phenomenon called allelopathy. morphine. spicy foods. Plants that produce cyanogenic glycosides also produce enzymes that convert these compounds into hydrogen cyanide. when herbivores feed on these tissues. Nicotine Figure 22. Caffeine is an alkaloid found in plants such as coffee (Coffea arabica). Figure 24. many defensive proteins . tea (Camellia sinensis). ultimately reducing enzyme function. Nicotine is an alkaloid that is produced in the roots of tobacco plants (Nicotiana tabacum) and transported to leaves where it is stored in vacuoles. These proteins are generally small and rich in the amino acid cysteine. also known as mustard oil glycosides. Unlike simple chemicals such as terpenoids.

the importance of beneficial microbes on plant health. pods. Hydrolytic enzymes are produced by some plants in response to pathogens and often accumulate in extracellular spaces where they degrade the cell walls of pathogenic fungi. cultural practices. and insect herbivores. but can be found in virtually all types of plant tissues including leaves. Castor beans (Ricinus communis) Digestive enzyme inhibitors are proteins that block the normal digestion and absorption of nutrients by vertebrate and invertebrate herbivores. Lysozymes are hydrolytic enzymes that are capable of degrading bacterial cell walls. however. and grasses. the reader is encouraged to investigate other aspects of plant defense including symbiotic relationships. including integrated pest management strategies. These defensins may inhibit preexisting ion channels or form new membrane pores that disrupt cellular ion balance. having an average lethal dose of only 0. and floral tissues. Defensins are best characterized in seeds. and genetic engineering. a class of polymers similar to cellulose that is present in the cell walls of many oomycetes (water molds). Protease inhibitors are typically produced in response to herbivore attack and inhibit digestive enzymes including trypsin and chymotrypsin. but they appear to act upon molecular aestivum) targets in the plasma membrane of pathogens. solanaceous plants. They occur widely in nature but have been well studied in legumes. Chitinases are enzymes that catalyze the degradation of chitin. Alpha-amylase inhibitors are proteins commonly found in legumes that bind to amylase enzymes and inhibit starch digestion. nematodes. fruit. Herbivore feeding often triggers a series of molecular signaling events that induce systemic production of these compounds in distal tissues that contribute to the protection of undamaged plant parts from subsequent attacks by a wide range of herbivore pests. and the impact of environmental conditions on plant disease. However. Wheat (Triticum and bacteria are still being characterized. biological control. Lectins are non-enzymatic proteins and glycoproteins that bind to carbohydrates and exhibit a wide range of functions including disruption of digestion in insects and agglutination of blood cells in vertebrates. Ricin is a highly potent toxin. They exhibit a wide range of biological activities that serve to inhibit the growth of many fungi and bacteria. bark. defensive proteins and enzymes effectively inhibit fungi.2 milligrams in humans. Ricin is a powerful toxin produced in castor beans (Ricinus communis). Figure 28.are only made in significant quantities after a pathogen or pest has attacked the plant. Some defensins also inhibit digestive proteins in herbivores. Additional Plant Defenses The mechanisms discussed in this article represent a broad overview of plant defense responses. a polymer with a backbone similar to cellulose that is present in the cell walls of true fungi. Once activated. One may also want to learn more about how humans influence disease resistance. tubers. bacteria. . Defensins are small cysteine-rich proteins that display broad anti-microbial activity and were first isolated from the endosperm of barley (Hordeum vulgare) and wheat (Triticum aestivum). They are widely distributed and may be present in most plants. Glucanases are enzymes that catalyze the degradation of glycosidic linkages in glucans. The precise mechanisms employed by plant defensins to inhibit fungi Figure 27. and transgenic plants expressing high levels of these enzymes exhibit increased resistance to a wide range of both foliar and root pathogens. In vitro analysis has verified the anti-fungal properties of these compounds. It combines a lectin molecule with an N-glycoside hydrolase that enters animal cells and inhibits protein synthesis. roots.


The deduced translation product shows high identity to Zea mays flavonoidglucosyltransferase. 2000)) catalyzes dehydration of p-hydroxyphenylacetaldehyde oxime to p-hydroxyphenylacetonitrile and C-hydroxylation of phydroxyphenylacetonitrile to p-hydroxymandelonitrile. the same products released from dhurrin upon cell disruption as a result of pest or herbivore attack (Bak et al. generating N. 1997). When both the two multifunctional sorghum cytochrome P450 enzymes CYP79A1 and CYP71E1 are constitutively expressed in tobacco and Arabidopsis p-hydroxymandelonitrile is formed. 1999). Sequential dehydration and decarboxylation reactions on this labile intermediate. 2000). Expression of CYP79A1 in Arabidopsis thaliana results in the production of high levels of the tyrosine-derived glucosinolate phydroxybenzylglucosinolate. p-Hydroxyphenylacetaldoxime is subsequently converted to p-hydroxymandelonitrile by a second multifunctional P450 monooxygenase (P450ox) (Schuler. 1999. In vitro reconstitution of the entire dhurrin biosynthetic pathway from tyrosine has been accomplished by the insertion of CYP79 (tyrosine N-hydroxylase). which is not a natural constituent of Arabidopsis (Bak et al. Note that conversion of an amino acid to an aldoxime by N-hydroxylation and decarboxylation. It is hypothesized that the glucosinolate pathway evolved from the cyanogenic glucoside pathway (Bak et al. CYP71E1 and UDP-glucose:p-hydroxymandelonitrile-Oglucosyltransferase (Tattersall et al. The accumulation of dhurrin in the transgenic plants confers resistance to . The final step in the biosynthesis of the cyanogenic glucoside dhurrin in sorghum is the transformation of the labile cyanohydrin into a stable storage form by O-glucosylation of (S)-p-hydroxymandelonitrile at the cyanohydrin function (Jones et al. Celenza. see Celenza (2001). probably proceeding nonenzymatically. 1996. The UDP-glucose:p-hydroxymandelonitrile-O-glucosyltransferase has been isolated and its cDNA cloned (Jones et al. 2001). 1999). The cytochrome P450 catalysing the conversion of tyrosine to p-hydroxyphenylacetaldoxime in the biosynthesis of the cyanogenic glucoside dhurrin in sorghum has been designated CYP79A1 (Bak et al. is also the first step in glucosinolate biosynthesis (see: Glucosinolates under Sulfate uptake and assimilation). Dhurrin (derived from the amino acid tyrosine) is the major cyanogenic glucoside in Sorghum bicolor (sorghum).Secondary products derived from aromatic amino acids Dhurrin synthesis For an early review of plant cyanogenic glucosides. 2001). Petersen et al.N-dihydroxytyrosine. P450ox. 1999). It may play a role in defense against herbivore attack. generate p-hydroxyphenylacetaldehyde oxime. P450ox (now known as CYP71E1 (Bak et al. see Poulton (1980). 1997). 1998. The entire pathway for synthesis of the tyrosine-derived cyanogenic glucoside dhurrin has been transferred from sorghum to Arabidopsis thaliana by expressing CYP79A1. and NADPH-P450 oxidoreductase in lipid micelles in the presence of uridine diphosphate glucose glucosyltransferase (Kahn et al. This provides further evidence that the enzymes of the glucosinolate pathway in Arabidopsis must have low substrate specificity with respect to aldoxime substrates (Bak et al. Dhurrin is synthesized via a multifunctional P450TYR (CYP79) that catalyzes two initial hydroxylations on tyrosine. 2001). Kahn et al. and can represent 30% of the dry weight of shoot tips of sorghum seedlings. This compound is labile and dissociates into p-hydroxybenzaldehyde and hydrogen cyanide. For a more recent review. 1999).

the flea beetle Phyllotreta nemorum. In Triglochin maritima (seaside arrow grass) the two cyanogenic glucosides taxiphyllin and triglochinin are derived from tyrosine in a pathway that is very similar to that of the dhurrin biosynthesis pathway in sorghum. 2001). This clearly demonstrates the importance of cyanogenic glucosides in plant defense against insect herbivores. Triglochin maritima has two cytochrome P450 enzymes of the CYP79 family (CYP79E1 and CYP79E2) that are functionally identical to that of sorghum CYP79A1 . a natural pest of other members of the crucifer group (Tattersall et al.

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