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A herbivore is an
animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
anatomically and physiologically adapted to eating
plant material Vascular tissue is a complex conducting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. There ...
, for example
foliage A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ...
or
marine algae Marine primary production is the chemical synthesis in the ocean of organic compounds from atmospheric or dissolved carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it al ...
, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthparts adapted to rasping or grinding.
Horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million yea ...
s and other herbivores have wide flat
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, ...
that are adapted to grinding
grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns a ...
,
tree bark Bark is the outermost layers of stems and roots of woody plants. Plants with bark include trees, woody vines, and shrubs. Bark refers to all the tissues outside the vascular cambium and is a nontechnical term. It overlays the wood and consist ...
, and other tough plant material. A large percentage of herbivores have mutualistic
gut flora Gut microbiota, gut microbiome, or gut flora, are the microorganisms, including bacteria, archaea, fungi, and viruses that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the gut m ...
that help them digest
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
matter, which is more difficult to digest than animal prey. This flora is made up of
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wa ...
-digesting
protozoans Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Hist ...
or
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
.


Etymology

Herbivore is the anglicized form of a modern Latin coinage, ''herbivora'', cited in
Charles Lyell Sir Charles Lyell, 1st Baronet, (14 November 1797 – 22 February 1875) was a Scottish geologist who demonstrated the power of known natural causes in explaining the earth's history. He is best known as the author of ''Principles of Geolo ...
's 1830 ''
Principles of Geology ''Principles of Geology: Being an Attempt to Explain the Former Changes of the Earth's Surface, by Reference to Causes Now in Operation'' is a book by the Scottish geologist Charles Lyell that was first published in 3 volumes from 1830–1833. Ly ...
''.J.A. Simpson and E.S.C. Weiner, eds. (2000) ''The Oxford English Dictionary'', vol. 8, p. 155.
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Ow ...
employed the anglicized term in an 1854 work on fossil teeth and skeletons. ''Herbivora'' is derived from Latin ''herba'' 'small plant, herb'P.G.W. Glare, ed. (1990) ''The Oxford Latin Dictionary'', p. 791. and ''vora'', from ''vorare'' 'to eat, devour'.P.G.W. Glare, ed. (1990) ''The Oxford Latin Dictionary'', p. 2103.


Definition and related terms

Herbivory is a form of
consumption Consumption may refer to: *Resource consumption *Tuberculosis, an infectious disease, historically * Consumption (ecology), receipt of energy by consuming other organisms * Consumption (economics), the purchasing of newly produced goods for curren ...
in which an
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...
principally eats
autotroph An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Wo ...
sAbraham, Martin A. A. Sustainability Science and Engineering, Volume 1. page 123. Publisher: Elsevier 2006. such as
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
s,
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...
and photosynthesizing
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
. More generally, organisms that feed on
autotroph An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Wo ...
s in general are known as primary consumers. Herbivory is usually limited to animals that eat plants. Fungi, bacteria, and protists that feed on living plants are usually termed plant pathogens (plant diseases), while fungi and
microbes A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
that feed on dead plants are described as
saprotroph Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi ( ...
s. Flowering plants that obtain nutrition from other living plants are usually termed
parasitic plant A parasitic plant is a plant that derives some or all of its nutritional requirements from another living plant. They make up about 1% of angiosperms and are found in almost every biome. All parasitic plants develop a specialized organ called th ...
s. There is, however, no single exclusive and definitive ecological classification of consumption patterns; each textbook has its own variations on the theme.


Evolution of herbivory

The understanding of herbivory in geological time comes from three sources: fossilized plants, which may preserve evidence of defence (such as spines), or herbivory-related damage; the observation of plant debris in fossilised animal faeces; and the construction of herbivore mouthparts. Although herbivory was long thought to be a
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
phenomenon, fossils have shown that plants were being consumed by arthropods within less than 20 million years after the first land plants evolved. Insects fed on the spores of early Devonian plants, and the
Rhynie chert The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness (a Lagerstätte). It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is located ...
also provides evidence that organisms fed on plants using a "pierce and suck" technique. During the next 75 million years, plants evolved a range of more complex organs, such as roots and seeds. There is no evidence of any organism being fed upon until the middle-late Mississippian, . There was a gap of 50 to 100 million years between the time each organ evolved and the time organisms evolved to feed upon them; this may be due to the low levels of oxygen during this period, which may have suppressed evolution. Further than their arthropod status, the identity of these early herbivores is uncertain. Hole feeding and skeletonization are recorded in the early
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleo ...
, with surface fluid feeding evolving by the end of that period. Herbivory among four-limbed terrestrial vertebrates, the
tetrapods Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct therapsi ...
, developed in the
Late Carboniferous Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effe ...
(307–299 million years ago). Early tetrapods were large amphibious
piscivores A piscivore () is a carnivorous animal that eats primarily fish. The name ''piscivore'' is derived . Piscivore is equivalent to the Greek-derived word ichthyophage, both of which mean "fish eater". Fish were the diet of early tetrapod evolut ...
. While amphibians continued to feed on fish and insects, some reptiles began exploring two new food types, tetrapods (carnivory) and plants (herbivory). The entire
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
order
ornithischia Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...
was composed of herbivorous dinosaurs. Carnivory was a natural transition from insectivory for medium and large tetrapods, requiring minimal adaptation. In contrast, a complex set of adaptations was necessary for feeding on highly fibrous plant materials.
Arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s evolved herbivory in four phases, changing their approach to it in response to changing plant communities.Tetrapod herbivores made their first appearance in the fossil record of their jaws near the Permio-Carboniferous boundary, approximately 300 million years ago. The earliest evidence of their herbivory has been attributed to dental occlusion, the process in which teeth from the upper jaw come in contact with teeth in the lower jaw is present. The evolution of dental occlusion led to a drastic increase in plant food processing and provides evidence about feeding strategies based on tooth wear patterns. Examination of
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
frameworks of tooth and jaw morphologes has revealed that dental occlusion developed independently in several lineages tetrapod herbivores. This suggests that evolution and spread occurred simultaneously within various lineages.


Food chain

Herbivores form an important link in the food chain because they consume plants to digest the carbohydrates photosynthetically produced by a plant.
Carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other s ...
s in turn consume herbivores for the same reason, while
omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nu ...
s can obtain their nutrients from either plants or animals. Due to a herbivore's ability to survive solely on tough and fibrous plant matter, they are termed the primary consumers in the food cycle (chain). Herbivory, carnivory, and omnivory can be regarded as special cases of
consumer–resource interactions Consumer–resource interactions are the core motif of ecological food chains or food webs, and are an umbrella term for a variety of more specialized types of biological species interactions including prey-predator (see predation), host-parasite ( ...
.


Feeding strategies

Two herbivore feeding strategies are
grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to roam around and consume wild vegetations in order to convert the otherwise indigestible (by human gut) cellulose within grass and ot ...
(e.g. cows) and
browsing Browsing is a kind of orienting strategy. It is supposed to identify something of relevance for the browsing organism. When used about human beings it is a metaphor taken from the animal kingdom. It is used, for example, about people browsing o ...
(e.g. moose). For a terrestrial mammal to be called a grazer, at least 90% of the forage has to be grass, and for a browser at least 90% tree leaves and twigs. An intermediate feeding strategy is called "mixed-feeding". In their daily need to take up energy from forage, herbivores of different body mass may be selective in choosing their food. "Selective" means that herbivores may choose their forage source depending on, e.g., season or food availability, but also that they may choose high quality (and consequently highly nutritious) forage before lower quality. The latter especially is determined by the body mass of the herbivore, with small herbivores selecting for high-quality forage, and with increasing body mass animals are less selective. Several theories attempt to explain and quantify the relationship between animals and their food, such as
Kleiber's law Kleiber's law, named after Max Kleiber for his biology work in the early 1930s, is the observation that, for the vast majority of animals, an animal's metabolic rate scales to the power of the animal's mass. Symbolically: if is the animal's meta ...
, Holling's disk equation and the marginal value theorem (see below). Kleiber's law describes the relationship between an animal's size and its feeding strategy, saying that larger animals need to eat less food per unit weight than smaller animals. Kleiber's law states that the metabolic rate (q0) of an animal is the mass of the animal (M) raised to the 3/4 power: q0=M3/4 Therefore, the mass of the animal increases at a faster rate than the metabolic rate. Herbivores employ numerous types of feeding strategies. Many herbivores do not fall into one specific feeding strategy, but employ several strategies and eat a variety of plant parts. Optimal Foraging Theory is a model for predicting animal behavior while looking for food or other resources, such as shelter or water. This model assesses both individual movement, such as animal behavior while looking for food, and distribution within a habitat, such as dynamics at the population and community level. For example, the model would be used to look at the browsing behavior of a deer while looking for food, as well as that deer's specific location and movement within the forested habitat and its interaction with other deer while in that habitat. This model has been criticized as circular and untestable. Critics have pointed out that its proponents use examples that fit the theory, but do not use the model when it does not fit the reality. Other critics point out that animals do not have the ability to assess and maximize their potential gains, therefore the optimal foraging theory is irrelevant and derived to explain trends that do not exist in nature. Holling's disk equation models the efficiency at which predators consume prey. The model predicts that as the number of prey increases, the amount of time predators spend handling prey also increases, and therefore the efficiency of the predator decreases. In 1959, S. Holling proposed an equation to model the rate of return for an optimal diet: Rate (R )=Energy gained in foraging (Ef)/(time searching (Ts) + time handling (Th))
R=Ef/(Ts + Th)
Where s=cost of search per unit time f=rate of encounter with items, h=handling time, e=energy gained per encounter
In effect, this would indicate that a herbivore in a dense forest would spend more time handling (eating) the vegetation because there was so much vegetation around than a herbivore in a sparse forest, who could easily browse through the forest vegetation. According to the Holling's disk equation, a herbivore in the sparse forest would be more efficient at eating than the herbivore in the dense forest. The
marginal value theorem The marginal value theorem (MVT) is an optimality model that usually describes the behavior of an optimally foraging individual in a system where resources (often food) are located in discrete patches separated by areas with no resources. Due to th ...
describes the balance between eating all the food in a patch for immediate energy, or moving to a new patch and leaving the plants in the first patch to regenerate for future use. The theory predicts that absent complicating factors, an animal should leave a resource patch when the rate of payoff (amount of food) falls below the average rate of payoff for the entire area. According to this theory, an animal should move to a new patch of food when the patch they are currently feeding on requires more energy to obtain food than an average patch. Within this theory, two subsequent parameters emerge, the Giving Up Density (GUD) and the Giving Up Time (GUT). The Giving Up Density (GUD) quantifies the amount of food that remains in a patch when a forager moves to a new patch. The Giving Up Time (GUT) is used when an animal continuously assesses the patch quality.


Plant-herbivore interactions

Interactions between plants and herbivores can play a prevalent role in
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...
dynamics such
community A community is a social unit (a group of living things) with commonality such as place, norms, religion, values, customs, or identity. Communities may share a sense of place situated in a given geographical area (e.g. a country, village, t ...
structure and functional processes. Plant diversity and
distribution Distribution may refer to: Mathematics * Distribution (mathematics), generalized functions used to formulate solutions of partial differential equations *Probability distribution, the probability of a particular value or value range of a vari ...
is often driven by herbivory, and it is likely that trade-offs between plant
competitiveness In economics, competition is a scenario where different economic firmsThis article follows the general economic convention of referring to all actors as firms; examples in include individuals and brands or divisions within the same (legal) firm ...
and defensiveness, and between colonization and mortality allow for coexistence between species in the presence of herbivores. However, the effects of herbivory on plant diversity and richness is variable. For example, increased abundance of herbivores such as deer decrease plant diversity and
species richness Species richness is the number of different species represented in an ecological community, landscape or region. Species richness is simply a count of species, and it does not take into account the abundances of the species or their relative ab ...
, while other large mammalian herbivores like bison control dominant species which allows other species to flourish.  Plant-herbivore interactions can also operate so that plant communities mediate herbivore communities. Plant communities that are more diverse typically sustain greater herbivore richness by providing a greater and more diverse set of resources.
Coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
and
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
correlation between herbivores and plants are important aspects of the influence of herbivore and plant interactions on communities and ecosystem functioning, especially in regard to herbivorous insects. This is apparent in the
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
s plants develop to
tolerate Toleration is the allowing, permitting, or acceptance of an action, idea, object, or person which one dislikes or disagrees with. Political scientist Andrew R. Murphy explains that "We can improve our understanding by defining "toleration" as a ...
and/or defend from insect herbivory and the responses of herbivores to overcome these adaptations. The evolution of antagonistic and mutualistic plant-herbivore interactions are not mutually exclusive and may co-occur. Plant phylogeny has been found to facilitate the colonization and community assembly of herbivores, and there is evidence of phylogenetic linkage between plant
beta diversity Beta (, ; uppercase , lowercase , or cursive ; grc, βῆτα, bē̂ta or ell, βήτα, víta) is the second letter of the Greek alphabet. In the system of Greek numerals, it has a value of 2. In Modern Greek, it represents the voiced labiod ...
and phylogenetic
beta diversity Beta (, ; uppercase , lowercase , or cursive ; grc, βῆτα, bē̂ta or ell, βήτα, víta) is the second letter of the Greek alphabet. In the system of Greek numerals, it has a value of 2. In Modern Greek, it represents the voiced labiod ...
of insect
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
s such as
butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises ...
. These types of eco-evolutionary feedbacks between plants and herbivores are likely the main driving force behind plant and herbivore diversity. Abiotic factors such as
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
and biogeographical features also impact plant-herbivore communities and interactions. For example, in temperate freshwater wetlands herbivorous waterfowl communities change according to season, with species that eat above-ground vegetation being abundant during summer, and species that forage below-ground being present in winter months. These seasonal herbivore communities differ in both their assemblage and functions within the wetland ecosystem. Such differences in herbivore modalities can potentially lead to trade-offs that influence species traits and may lead to additive effects on
community A community is a social unit (a group of living things) with commonality such as place, norms, religion, values, customs, or identity. Communities may share a sense of place situated in a given geographical area (e.g. a country, village, t ...
composition and ecosystem functioning. Seasonal changes and environmental gradients such as elevation and
latitude In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north ...
often affect the
palatability Palatability (or palatableness) is the hedonic reward (i.e., pleasure Pleasure refers to experience that feels good, that involves the enjoyment of something. It contrasts with pain or suffering, which are forms of feeling bad. It is closely ...
of plants which in turn influences herbivore community assemblages and vice versa. Examples include a decrease in abundance of leaf-chewing larvae in the fall when hardwood leaf palatability decreases due to increased
tannin Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' (from Anglo-Norman ''tanner'' ...
levels which results in a decline of arthropod
species richness Species richness is the number of different species represented in an ecological community, landscape or region. Species richness is simply a count of species, and it does not take into account the abundances of the species or their relative ab ...
, and increased palatability of plant communities at higher elevations where
grasshopper Grasshoppers are a group of insects belonging to the suborder Caelifera. They are among what is possibly the most ancient living group of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago. Grasshopp ...
s abundances are lower. Climatic stressors such as
ocean acidification Ocean acidification is the reduction in the pH value of the Earth’s ocean. Between 1751 and 2021, the average pH value of the ocean surface has decreased from approximately 8.25 to 8.14. The root cause of ocean acidification is carbon dioxid ...
can lead to responses in plant-herbivore interactions in relation to palatability as well.


Herbivore offense

The myriad defenses displayed by plants means that their herbivores need a variety of skills to overcome these defenses and obtain food. These allow herbivores to increase their feeding and use of a host plant. Herbivores have three primary strategies for dealing with plant defenses: choice, herbivore modification, and plant modification. Feeding choice involves which plants a herbivore chooses to consume. It has been suggested that many herbivores feed on a variety of plants to balance their nutrient uptake and to avoid consuming too much of any one type of defensive chemical. This involves a tradeoff however, between foraging on many plant species to avoid toxins or specializing on one type of plant that can be detoxified. Herbivore modification is when various adaptations to body or digestive systems of the herbivore allow them to overcome plant defenses. This might include detoxifying secondary metabolites, sequestering toxins unaltered, or avoiding toxins, such as through the production of large amounts of saliva to reduce effectiveness of defenses. Herbivores may also utilize symbionts to evade plant defenses. For example, some aphids use bacteria in their gut to provide essential amino acids lacking in their sap diet. Plant modification occurs when herbivores manipulate their plant prey to increase feeding. For example, some caterpillars roll leaves to reduce the effectiveness of plant defenses activated by sunlight.


Plant defense

A plant defense is a trait that increases plant fitness when faced with herbivory. This is measured relative to another plant that lacks the defensive trait. Plant defenses increase survival and/or reproduction (fitness) of plants under pressure of predation from herbivores. Defense can be divided into two main categories, tolerance and resistance. Tolerance is the ability of a plant to withstand damage without a reduction in fitness. This can occur by diverting herbivory to non-essential plant parts, resource allocation, compensatory growth, or by rapid regrowth and recovery from herbivory. Resistance refers to the ability of a plant to reduce the amount of damage it receives from herbivores. This can occur via avoidance in space or time, physical defenses, or chemical defenses. Defenses can either be constitutive, always present in the plant, or induced, produced or translocated by the plant following damage or stress. Physical, or mechanical, defenses are barriers or structures designed to deter herbivores or reduce intake rates, lowering overall herbivory. Thorns such as those found on roses or acacia trees are one example, as are the spines on a cactus. Smaller hairs known as
trichomes Trichomes (); ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a plant ...
may cover leaves or stems and are especially effective against invertebrate herbivores. In addition, some plants have
waxes Waxes are a diverse class of organic compounds that are lipophilic, malleable solids near ambient temperatures. They include higher alkanes and lipids, typically with melting points above about 40 °C (104 °F), melting to give low ...
or resins that alter their texture, making them difficult to eat. Also the incorporation of silica into cell walls is analogous to that of the role of lignin in that it is a compression-resistant structural component of cell walls; so that plants with their cell walls impregnated with silica are thereby afforded a measure of protection against herbivory. Chemical defenses are
secondary metabolites Secondary metabolites, also called specialised metabolites, toxins, secondary products, or natural products, are organic compounds produced by any lifeform, e.g. bacteria, fungi, animals, or plants, which are not directly involved in the nor ...
produced by the plant that deter herbivory. There are a wide variety of these in nature and a single plant can have hundreds of different chemical defenses. Chemical defenses can be divided into two main groups, carbon-based defenses and nitrogen-based defenses. #Carbon-based defenses include
terpenes Terpenes () are a class of natural products consisting of compounds with the formula (C5H8)n for n > 1. Comprising more than 30,000 compounds, these unsaturated hydrocarbons are produced predominantly by plants, particularly conifers. Terpenes ar ...
and phenolics. Terpenes are derived from 5-carbon isoprene units and comprise essential oils, carotenoids, resins, and latex. They can have several functions that disrupt herbivores such as inhibiting adenosine triphosphate (ATP) formation, molting
hormones A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are required ...
, or the nervous system. Phenolics combine an aromatic carbon ring with a hydroxyl group. There are several different phenolics such as lignins, which are found in cell walls and are very indigestible except for specialized microorganisms; tannins, which have a bitter taste and bind to proteins making them indigestible; and furanocumerins, which produce free radicals disrupting DNA, protein, and lipids, and can cause skin irritation. #Nitrogen-based defenses are synthesized from amino acids and primarily come in the form of
alkaloids Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar ...
and cyanogens. Alkaloids include commonly recognized substances such as
caffeine Caffeine is a central nervous system (CNS) stimulant of the methylxanthine chemical classification, class. It is mainly recreational drug use, used recreationally as a Nootropic, cognitive enhancer, increasing alertness and attentional perfor ...
,
nicotine Nicotine is a naturally produced alkaloid in the nightshade family of plants (most predominantly in tobacco and '' Duboisia hopwoodii'') and is widely used recreationally as a stimulant and anxiolytic. As a pharmaceutical drug, it is use ...
, and
morphine Morphine is a strong opiate that is found naturally in opium, a dark brown resin in poppies ('' Papaver somniferum''). It is mainly used as a pain medication, and is also commonly used recreationally, or to make other illicit opioids. T ...
. These compounds are often bitter and can inhibit DNA or RNA synthesis or block nervous system signal transmission. Cyanogens get their name from the
cyanide Cyanide is a naturally occurring, rapidly acting, toxic chemical that can exist in many different forms. In chemistry, a cyanide () is a chemical compound that contains a functional group. This group, known as the cyano group, consists of ...
stored within their tissues. This is released when the plant is damaged and inhibits cellular respiration and electron transport. Plants have also changed features that enhance the probability of attracting natural enemies to herbivores. Some emit semiochemicals, odors that attract natural enemies, while others provide food and housing to maintain the natural enemies' presence, e.g.
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s that reduce herbivory. A given plant species often has many types of defensive mechanisms, mechanical or chemical, constitutive or induced, which allow it to escape from herbivores.


Predator–Prey Theory

According to the theory of
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
–prey interactions, the relationship between herbivores and plants is cyclic. When prey (plants) are numerous their predators (herbivores) increase in numbers, reducing the prey population, which in turn causes predator number to decline. The prey population eventually recovers, starting a new cycle. This suggests that the population of the herbivore fluctuates around the carrying capacity of the food source, in this case, the plant. Several factors play into these fluctuating populations and help stabilize predator-prey dynamics. For example, spatial heterogeneity is maintained, which means there will always be pockets of plants not found by herbivores. This stabilizing dynamic plays an especially important role for specialist herbivores that feed on one species of plant and prevents these specialists from wiping out their food source. Prey defenses also help stabilize predator-prey dynamics, and for more information on these relationships see the section on Plant Defenses. Eating a second prey type helps herbivores' populations stabilize. Alternating between two or more plant types provides population stability for the herbivore, while the populations of the plants oscillate. This plays an important role for generalist herbivores that eat a variety of plants. Keystone herbivores keep vegetation populations in check and allow for a greater diversity of both herbivores and plants. When an invasive herbivore or plant enters the system, the balance is thrown off and the diversity can collapse to a monotaxon system. The back and forth relationship of plant defense and herbivore offense drives
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
between plants and herbivores, resulting in a "coevolutionary arms race". The escape and radiation mechanisms for coevolution, presents the idea that adaptations in herbivores and their host plants, has been the driving force behind
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution withi ...
.Thompson, J. 1999. What we know and do not know about coevolution: insect herbivores and plants as a test case. Pages 7–30 in H. Olff, V. K. Brown, R. H. Drent, and British Ecological Society Symposium 1997 (Corporate Author), editors. Herbivores: between plants and predators. Blackwell Science, London, UK.


Mutualism

While much of the interaction of herbivory and plant defense is negative, with one individual reducing the fitness of the other, some is beneficial. This beneficial herbivory takes the form of mutualisms in which both partners benefit in some way from the interaction.
Seed dispersal In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vector ...
by herbivores and
pollination Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...
are two forms of mutualistic herbivory in which the herbivore receives a food resource and the plant is aided in reproduction. Plants can also be indirectly affected by herbivores through nutrient recycling, with plants benefiting from herbivores when nutrients are recycled very efficiently. Another form of plant-herbivore mutualism is physical changes to the environment and/or plant community structure by herbivores which serve as
ecosystem engineer An ecosystem engineer is any species that creates, significantly modifies, maintains or destroys a habitat. These organisms can have a large impact on species richness and landscape-level heterogeneity of an area. As a result, ecosystem engin ...
s, such as wallowing by bison. Swans form a mutual relationship with the plant species that they
forage Forage is a plant material (mainly plant leaves and stems) eaten by grazing livestock. Historically, the term ''forage'' has meant only plants eaten by the animals directly as pasture, crop residue, or immature cereal crops, but it is also use ...
by digging and disturbing the sediment which removes competing plants and subsequently allows colonization of other plant species.


Impacts


Trophic Cascades and Environmental Degradation

When herbivores are affected by
trophic cascade Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to reduce t ...
s, plant communities can be indirectly affected. Often these effects are felt when predator populations decline and herbivore populations are no longer limited, which leads to intense herbivore foraging which can suppress plant communities. With the size of herbivores having an effect on the amount of energy intake that is needed, larger herbivores need to forage on higher quality or more plants to gain the optimal amount of nutrients and energy compared to smaller herbivores.
Environmental degradation Environmental degradation is the deterioration of the environment through depletion of resources such as quality of air, water and soil; the destruction of ecosystems; habitat destruction; the extinction of wildlife; and pollution. It is defin ...
from
white-tailed deer The white-tailed deer (''Odocoileus virginianus''), also known as the whitetail or Virginia deer, is a medium-sized deer native to North America, Central America, and South America as far south as Peru and Bolivia. It has also been introduced t ...
(''Odocoileus virginianus'') in the US alone has the potential to both change vegetative communities through over-browsing and cost
forest restoration Forest restoration is defined as “actions to re-instate ecological processes, which accelerate recovery of forest structure, ecological functioning and biodiversity levels towards those typical of climax forest” i.e. the end-stage of natural ...
projects upwards of $750 million annually. Another example of a trophic cascade involved plant-herbivore interactions are
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of Colony (biology), colonies of coral polyp (zoology), polyps held together by calcium carbonate. Most coral reefs are built from stony corals, wh ...
ecosystems.
Herbivorous fish A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
and marine animals are important
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...
and seaweed grazers, and in the absence of plant-eating fish, corals are outcompeted and seaweeds deprive corals of sunlight.


Economic Impacts

Agricultural crop damage by the same species totals approximately $100 million every year. Insect crop damages also contribute largely to annual crop losses in the U.S.An Integrated Approach To Deer Damage Control Publication No. 809 West Virginia Division of Natural Resources Cooperative Extension Service, Wildlife Resources Section West Virginia University, Law Enforcement Section Center for Extension and Continuing Education, March 1999 Herbivores also affect economics through the revenue generated by hunting and ecotourism. For example, the hunting of herbivorous game species such as white-tailed deer, cottontail rabbits, antelope, and elk in the U.S. contributes greatly to the billion-dollar annually, hunting industry.
Ecotourism Ecotourism is a form of tourism involving responsible travel (using sustainable transport) to natural areas, conserving the environment, and improving the well-being of the local people. Its purpose may be to educate the traveler, to provide fund ...
is a major source of revenue, particularly in Africa, where many large mammalian herbivores such as elephants, zebras, and giraffes help to bring in the equivalent of millions of US dollars to various nations annually.


See also

* Consumer-resource systems *
List of feeding behaviours Feeding is the process by which organisms, typically animals, obtain food. Terminology often uses either the suffixes -vore, -vory, or -vorous from Latin ''vorare'', meaning "to devour", or -phage, -phagy, or -phagous from Greek φαγε ...
*
List of herbivorous animals Herbivores are animals that eat plants. Herbivory is a form of consumption in which a heterotrophic organism consumes other organisms, principally autotrophsCampbell, N. A. (1996) ''Biology'' (4th edition) Benjamin Cummings, New York such as pl ...
*
Plant-based diet A plant-based diet is a diet consisting mostly or entirely of plant-based foods. Plant-based diets encompass a wide range of dietary patterns that contain low amounts of animal products and high amounts of plant products such as vegetables, fru ...
*
Productivity (ecology) In ecology, the term productivity refers to the rate of generation of biomass in an ecosystem, usually expressed in units of mass per volume (unit surface) per unit of time, such as grams per square metre per day (g m−2 d−1). The unit of mass ...
*
Seed predation Seed predation, often referred to as granivory, is a type of plant-animal interaction in which granivores (seed predators) feed on the seeds of plants as a main or exclusive food source,Hulme, P.E. and Benkman, C.W. (2002) "Granivory", pp. 13 ...
* Tritrophic interactions in plant defense * Herbivore men


References


Further reading

* Bob Strauss, 2008
Herbivorous Dinosaurs
The New York Times ''The New York Times'' (''the Times'', ''NYT'', or the Gray Lady) is a daily newspaper based in New York City with a worldwide readership reported in 2020 to comprise a declining 840,000 paid print subscribers, and a growing 6 million paid ...
*Danell, K., R. Bergström, P. Duncan, J. Pastor (Editors)(2006) ''Large herbivore ecology, ecosystem dynamics and conservation'' Cambridge, UK : Cambridge University Press. 506 p.  *Crawley, M. J. (1983) ''Herbivory : the dynamics of animal-plant interactions'' Oxford : Blackwell Scientific. 437 p.  *Olff, H., V.K. Brown, R.H. Drent (editors) (1999) ''Herbivores : between plants and predators'' Oxford ; Malden, Ma. : Blackwell Science. 639 p. 


External links


Herbivore information resource websiteHerbivore defense in ''Lindera benzoin''website of the herbivory lab at Cornell University
{{Authority control * Ecology terminology Animals by eating behaviors