Escape and radiate coevolution

Last updated
Garter snake is an example of escape and radiate coevolution. Coast Garter Snake.jpg
Garter snake is an example of escape and radiate coevolution.

Escape and radiate coevolution is a hypothesis proposing that a coevolutionary 'arms-race' between primary producers and their consumers contributes to the diversification of species by accelerating speciation rates. The hypothesized process involves the evolution of novel defenses in the host, allowing it to "escape" and then "radiate" into differing species.

Contents

History

This hypothesis originated in a 1964 paper by Paul Ehrlich and Peter Raven, "Butterflies and plants: a study in coevolution". [1] While this paper outlined the concept, the actual term "escape and radiate" was not actually coined until 1989 by John N. Thompson. [2] [ citation needed ] The theory has been highly influential in chemical ecology and plant evolutionary ecology, but remains controversial due to the difficulty of collecting decisive evidence [3] as well as uncertainty about the mechanisms linking ecological 'escape' with evolutionary diversification. [4]

Theory

Escape

A variety of defense mechanisms can lead to ecological escape from predators. Plants use chemical defenses in the form of secondary metabolites or allelochemicals. These allelochemicals inhibit the growth, behavior, and health of herbivores, allowing plants to escape. [5] An example of a plant allelochemical are alkaloids that can inhibit protein synthesis in herbivores. Other forms of plant defense include mechanical defenses such as thigmonasty movements which have the plant leaves close in response to tactile stimulation. Indirect mechanisms plant include shedding of plant leaves so less leaves are available which deters herbivores, growth in locations in that are difficult to reach, and even mimicry. For organisms other than plants, examples of defense mechanisms allowing for escape include camouflage, aposematism, [6] heightened senses and physical capabilities, and even defensive behaviors such as feigning death. An example of an organism using one of these defense mechanisms is the granular poison frog which defends itself through aposematism. It is important to understand that in order for escape and radiate coevolution to occur, it is necessary that the developed defense is novel rather than previously established.

Induced defense stemming from adaptive phenotypic plasticity may help a plant defend itself against multiple enemies. [7] Phenotypic plasticity occurs when an organism undergoes an environmental change forcing a change altering its behavior, physiology, etc. These induced defenses allow for an organism to escape.

Radiation

Radiation is the evolutionary process of diversification of a single species into multiple forms. It includes the physiological and ecological diversity within a rapidly multiplying lineage. [8] There are many types of radiation including adaptive, concordant, and discordant radiation however escape and radiate coevolution does not always follow those specific types.

Ehrlich and Raven's original paper did not clearly answer why ecological escape leads to increased diversification, however several explanations have been proposed. [4] [9] Once a novel defense has been acquired, the attacking organism which had evolved adaptations that allowed it to predate is now up against a new defense that it has not yet been evolved to encounter. This gives the defending organism the advantage, and therefore time to rapidly multiply unopposed by the previously attacking organism. This ultimately leads to the physiological and ecological diversity within the rapidly multiplying lineage, hence radiation.

Importance

Ehrlich and Raven's paper was highly influential on a generation of biologists and contributed to the explosion of research on plant-insect interactions and chemical ecology. The theory of escape and radiate coevolution purports to explain why we see such vast biological diversity on earth. After the organism escapes, it then radiates into multiple species, and spreads geographically. Evidence of escape and radiate coevolution can be seen through the starburst effect in plant and herbivore clades. [10] When analyzing clades of predator-prey associations, although it varies, the starburst effect is a good indicator that escape and radiate coevolution may be occurring. Eventually this cycle must come to an end because adaptations that entail costs (such as allocation of resources, or vulnerability to other predators) at some point outweigh their benefits. [11]

Escape and radiate coevolution may support parallel cladogenesis, wherein plant and herbivore phylogenies might match with ancestral insects exploiting ancestral plants. This is significant because it allows researchers to hypothesize about the relationships between ancestral organisms. [12] Unfortunately, there have not yet been any known examples specifically involving escape and radiate coevolution being used for hypothesizing ancestral relationships.

Many times the organism that has "escaped" continuously undergoes selective pressure because the predator it has escaped from evolves to create another adaptation in response, causing the process to continue. These "offensive" traits developed by predators range widely. For example, herbivores can develop an adaptation that allows for improved detoxification which allow to overcome plant defenses, thus causing escape and radiate coevolution to continue. Often the term "evolutionary arms race" is used to illustrate the idea that continuous evolution is needed to maintain the same relative fitness while the two species are coevolving. This idea also ties in with the Red Queen hypothesis. Counter adaptations among two organisms through escape and radiate coevolution is a major driving force behind diversity.

Escape and radiate coevolution produces much more biological variation than other evolutionary mechanisms. For instance, cospeciation is important for diversity amongst species that share a symbiotic relationship, however this does not create nearly as much diversity in comparison to reciprocal evolutionary change due to natural selection. [13] Evidence of rapid diversification following a novel adaptation is shown through the evolution of resin and latex canal tubes in 16 different lineages of plants. Plants with resin or latex canals can easily defend themselves against insect herbivores. When lineages of canal bearing plants are compared to the lineages of canal free plants, it is apparent that canal bearing plants are far more diverse, supporting escape and radiate coevolution. [14]

Examples

Plant-herbivore

Erlich and Raven's Classic butterfly model Monarch In May.jpg
Erlich and Raven's Classic butterfly model

The most popular examples of escape and radiate coevolution are of plant-herbivore associations. The most classic example is of butterflies and plants outlined in Ehrlich and Raven's original paper, "Butterflies and plants: a study in coevolution.". Erlich and Raven found in 1964 that hostplants for butterflies had a wide range of chemical defenses, allowing them to escape herbivory. Butterflies who developed novel counter detoxification mechanisms against the hostplants chemical defenses were able to utilize the hostplant resources. [1] The process of stepwise adaptation and counteradaptation among the butterflies and hostplants is continuous and creates vast diversity.

Tropical trees may also escape and defend themselves. Trees growing in high light were predicted to have few chemical defenses, but rapid synchronous leaf expansion and low leaf nutritional quality during expansion. Species growing in low light have high levels of different chemical defenses, poor nutritional quality and asynchronous leaf expansion. Depending on the level of light the trees were growing in influenced the type of defenses they obtained, either chemical or through leaf expansion. The trees exposed to less light developed various chemicals to defend themselves against herbivores, a defense not utilizing light. This study was significant because it illustrates the separation between defenses and their relationship with an organism escaping and radiating into other species. Development of novel defenses does not necessarily imply that escape is possible for a species of plant if herbivores are adapting at a faster rate. [15]

Common milkweed plant bears latex tubes to ward off herbivores. Asclepiascommon.JPG
Common milkweed plant bears latex tubes to ward off herbivores.

Milkweed plants contain latex-filled canals which deter insect herbivores. Milkweed latex not only gums up the mouth-parts of insects but is also toxic because it contains cardenolides which disrupt sodium and potassium levels by inhibiting the essential enzyme, Na+/K+‐ATPase. This has allowed for milkweeds to "escape" and become extremely diverse. There are over 100 different species of milkweeds which shows how diverse the plant is, with escape and radiate coevolution playing a very large role in creating such a high number of species. [16]

Fish-water flea

Key adaptations are adaptations that allow a group of organisms to diversify. Daphnia lumholtzi is a water flea that is able to form rigid head spines in response to chemicals released when fish are present. These phenotypically plastic traits serve as an induced defense against these predators. A study showed that Daphnia pulicaria is competitively superior to D. lumholtzi in the absence of predators. However, in the presence of fish predation the invasive species formed its defenses and became the dominant water flea in the region. This switch in dominance suggests that the induced defense against fish predation could represent a key adaptation for the invasion success of D. lumholtzi. A defensive trait that qualifies as a key adaptation is most likely an example of escape and radiate coevolution. [17]

Bacteria-phage

The theory can be applied at the microscopic level such as to bacteria-phage relationships. Bacteria were able to diversify and escape through resistance to phages. The diversity among the hosts and parasites differed among the range of infection and resistance. The implication of this study to humans is its important to understanding the evolution of infectious organisms, and preventing diseases. [18]

Related Research Articles

Herbivore Organism that eats mostly or exclusively plant material

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 mouthparts adapted to rasping or grinding. Horses and other herbivores have wide flat teeth that are adapted to grinding grass, tree bark, and other tough plant material.

<i>Erysimum</i> Genus of flowering plants

Erysimum, or wallflower, is a genus of flowering plants in the cabbage family, Brassicaceae. It includes more than 150 species, both popular garden plants and many wild forms. The genus Cheiranthus is sometimes included here in whole or in part. Erysimum has since the early 21st century been ascribed to a monogeneric cruciferous tribe, Erysimeae, characterised by sessile, stellate (star-shaped) and/or malpighiaceous (two-sided) trichomes, yellow to orange flowers and multiseeded siliques.

Coevolution Two or more species influencing each others evolution

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 as gene-culture coevolution.

<i>Asclepias</i> Genus of flowering plants

Asclepias is a genus of herbaceous, perennial, flowering plants known as milkweeds, named for their latex, a milky substance containing cardiac glycosides termed cardenolides, exuded where cells are damaged. Most species are toxic to humans and many other species, primarily due to the presence of cardenolides, although, as with many such plants, there are species that feed upon them and from them. Most notable are monarch butterflies, who use and require certain milkweeds as host plants for their larvae.

Chemical ecology is the study of chemically-mediated interactions between living organisms, and the effects of those interactions on the demography, behavior and ultimately evolution of the organisms involved. It is thus a vast and highly interdisciplinary field. Chemical ecologists seek to identify the specific molecules that function as signals mediating community or ecosystem processes and to understand the evolution of these signals. The substances that serve in such roles are typically small, readily-diffusible organic molecules, but can also include larger molecules and small peptides.

An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. Radiations may affect one clade or many, and be rapid or gradual; where they are rapid, and driven by a single lineage's adaptation to their environment, they are termed adaptive radiations.

A generalist species is able to thrive in a wide variety of environmental conditions and can make use of a variety of different resources. A specialist species can thrive only in a narrow range of environmental conditions or has a limited diet. Most organisms do not all fit neatly into either group, however. Some species are highly specialized, others less so, and some can tolerate many different environments. In other words, there is a continuum from highly specialized to broadly generalist species.

Plant defense against herbivory Plants defenses against being eaten

Plant defense against herbivory or host-plant resistance (HPR) describes a range of adaptations evolved by plants which improve their survival and reproduction by reducing the impact of herbivores. Plants can sense being touched, and they can use several strategies to defend against damage caused by herbivores. Many plants produce secondary metabolites, known as allelochemicals, that influence the behavior, growth, or survival of herbivores. These chemical defenses can act as repellents or toxins to herbivores, or reduce plant digestibility. Another defensive strategy of plants is changing their attractiveness. To prevent overconsumption by large herbivores, plants alter their appearance by changing their size or quality, overall decreasing their consumption rate.

Herbivores are dependent on plants for food, and have coevolved mechanisms to obtain this food despite the evolution of a diverse arsenal of plant defenses against herbivory. Herbivore adaptations to plant defense have been likened to "offensive traits" and consist of those traits that allow for increased feeding and use of a host. Plants, on the other hand, protect their resources for use in growth and reproduction, by limiting the ability of herbivores to eat them. Relationships between herbivores and their host plants often results in reciprocal evolutionary change. When a herbivore eats a plant it selects for plants that can mount a defensive response, whether the response is incorporated biochemically or physically, or induced as a counterattack. In cases where this relationship demonstrates "specificity", and "reciprocity", the species are thought to have coevolved. 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. The coevolution that occurs between plants and herbivores that ultimately results in the speciation of both can be further explained by the Red Queen hypothesis. This hypothesis states that competitive success and failure evolve back and forth through organizational learning. The act of an organism facing competition with another organism ultimately leads to an increase in the organism's performance due to selection. This increase in competitive success then forces the competing organism to increase its performance through selection as well, thus creating an "arms race" between the two species. Herbivores evolve due to plant defenses because plants must increase their competitive performance first due to herbivore competitive success.

<i>Tetraopes tetrophthalmus</i> Species of beetle

The red milkweed beetle is a beetle in the family Cerambycidae.

Myrmecophily Positive interspecies associations between ants and other organisms

Myrmecophily is the term applied to positive interspecies associations between ants and a variety of other organisms, such as plants, other arthropods, and fungi. Myrmecophily refers to mutualistic associations with ants, though in its more general use, the term may also refer to commensal or even parasitic interactions.

Cardenolide Chemical compound

A cardenolide is a type of steroid. Many plants contain derivatives, collectively known as cardenolides, including many in the form of cardenolide glycosides (cardenolides that contain structural groups derived from sugars). Cardenolide glycosides are often toxic; specifically, they are heart-arresting. Cardenolides are toxic to animals through inhibition of the enzyme Na+/K+‐ATPase, which is responsible for maintaining the sodium and potassium ion gradients across the cell membranes.

Chemical mimicry

Chemical mimicry is a type of biological mimicry, involving the use of chemicals to dupe an operator. A chemical mimic dupes an operator by showing an adaptive chemical resemblance to an object of its environment and as a consequence receives selective advantage. In all cases of chemical mimicry it has been found that the mimicking species is the only species to benefit from the reaction with either costs or no effect on the duped species. This is by adapting to produce chemicals that will cause a desirable behavioural reaction in the species being deceived and a selective advantage to the mimic. Chemical mimicry exists within many of the different forms of mimicry such as aggressive, protective, Batesian, and Müllerian mimicry and can involve a number of different senses. Mimicking semiochemicals, which cannot be seen, make up some of the most widely used forms of chemical mimicry and is therefore less apparent than more visual forms. As a result of this, this topic has been relatively neglected in research and literature. Two examples of organisms displaying chemical mimicry include the mimicking of Noctuid pheromones by bolas spiders in order to draw prey to the spider’s location and the duping of insects within their own nests by mimicking their odours in order to enter and hide within the nest undetected. It is important to note that in all forms of mimicry the mimicking organism is not conscious of the deceit used and does not act intentionally to trick other organisms.

Ecological fitting Biological process

Ecological fitting is "the process whereby organisms colonize and persist in novel environments, use novel resources or form novel associations with other species as a result of the suites of traits that they carry at the time they encounter the novel condition". It can be understood as a situation in which a species' interactions with its biotic and abiotic environment seem to indicate a history of coevolution, when in actuality the relevant traits evolved in response to a different set of biotic and abiotic conditions.

Chemical defense

Chemical defense is a life history strategy employed by many organisms to avoid consumption by producing toxic or repellent metabolites. The production of defensive chemicals occurs in plants, fungi, and bacteria, as well as invertebrate and vertebrate animals. The class of chemicals produced by organisms that are considered defensive may be considered in a strict sense to only apply to those aiding an organism in escaping herbivory or predation. However, the distinction between types of chemical interaction is subjective and defensive chemicals may also be considered to protect against reduced fitness by pests, parasites, and competitors. Many chemicals used for defensive purposes are secondary metabolites derived from primary metabolites which serve a physiological purpose in the organism. Secondary metabolites produced by plants are consumed and sequestered by a variety of arthropods and, in turn, toxins found in some amphibians, snakes, and even birds can be traced back to arthropod prey. There are a variety of special cases for considering mammalian antipredatory adaptations as chemical defenses as well.

Insects have a wide variety of predators, including birds, reptiles, amphibians, mammals, carnivorous plants, and other arthropods. The great majority (80–99.99%) of individuals born do not survive to reproductive age, with perhaps 50% of this mortality rate attributed to predation. In order to deal with this ongoing escapist battle, insects have evolved a wide range of defense mechanisms. The only restraint on these adaptations is that their cost, in terms of time and energy, does not exceed the benefit that they provide to the organism. The further that a feature tips the balance towards beneficial, the more likely that selection will act upon the trait, passing it down to further generations. The opposite also holds true; defenses that are too costly will have a little chance of being passed down. Examples of defenses that have withstood the test of time include hiding, escape by flight or running, and firmly holding ground to fight as well as producing chemicals and social structures that help prevent predation.

In evolutionary biology, a key innovation, also known as an adaptive breakthrough or key adaptation, is a novel phenotypic trait that allows subsequent radiation and success of a taxonomic group. Typically they bring new abilities that allows the taxa to rapidly diversify and invade niches that were not previously available. The phenomenon helps to explain how some taxa are much more diverse and have many more species than their sister taxa. The term was first used in 1949 by Alden H. Miller who defined it as "key adjustments in the morphological and physiological mechanism which are essential to the origin of new major groups", although a broader, contemporary definition holds that "a key innovation is an evolutionary change in individual traits that is causally linked to an increased diversification rate in the resulting clade".

Evolving digital ecological network

Evolving digital ecological networks are webs of interacting, self-replicating, and evolving computer programs that experience the same major ecological interactions as biological organisms. Despite being computational, these programs evolve quickly in an open-ended way, and starting from only one or two ancestral organisms, the formation of ecological networks can be observed in real-time by tracking interactions between the constantly evolving organism phenotypes. These phenotypes may be defined by combinations of logical computations that digital organisms perform and by expressed behaviors that have evolved. The types and outcomes of interactions between phenotypes are determined by task overlap for logic-defined phenotypes and by responses to encounters in the case of behavioral phenotypes. Biologists use these evolving networks to study active and fundamental topics within evolutionary ecology.

Tritrophic interactions in plant defense Ecological interactions

Tritrophic interactions in plant defense against herbivory describe the ecological impacts of three trophic levels on each other: the plant, the herbivore, and its natural enemies. They may also be called multitrophic interactions when further trophic levels, such as soil microbes, endophytes, or hyperparasitoids are considered. Tritrophic interactions join pollination and seed dispersal as vital biological functions which plants perform via cooperation with animals.

Calotropin Chemical compound

Calotropin is a toxic cardenolide found in plants in the family Asclepiadoideae. In extreme cases, calotropin poisoning can cause respiratory and cardiac failure. Accidental poisoning is common in livestock who have ingested milkweed. Calotropin is commonly stored as a defense mechanism by insects that eat milkweeds as their main food source.

References

  1. 1 2 Ehrlich, P. R., & Raven, P. H. (1964). Butterflies and plants: a study in coevolution. Evolution, 586-608.
  2. Thompson, 1989
  3. Suchan, Tomasz; Alvarez, Nadir (2015). "Fifty years after Ehrlich and Raven, is there support for plant–insect coevolution as a major driver of species diversification?". Entomologia Experimentalis et Applicata. 157 (1): 98–112. doi: 10.1111/eea.12348 . ISSN   1570-7458 . Retrieved 2020-10-18.
  4. 1 2 Janz, Niklas (2011). "Ehrlich and Raven Revisited: Mechanisms Underlying Codiversification of Plants and Enemies". Annual Review of Ecology, Evolution, and Systematics. 42 (1): 71–89. doi:10.1146/annurev-ecolsys-102710-145024 . Retrieved 2020-10-18.
  5. Haig, Terry. "Allelochemicals in plants." Allelopathy in Sustainable Agriculture and Forestry. Springer New York, 2008. 63-104.
  6. Mappes, Johanna, Nicola Marples, and John A. Endler. "The complex business of survival by aposematism." Trends in Ecology & Evolution 20.11 (2005): 598-603.
  7. Agrawal, Anurag A. "Induced plant defense: evolution of induction and adaptive phenotypic plasticity." Inducible plant defenses against pathogens and herbivores: biochemistry, ecology, and agriculture. American Phytopathological Society Press, St. Paul, MN (1999): 251-268.
  8. Schluter, Dolph. The ecology of adaptive radiation. Oxford University Press, 2000.
  9. Marquis, Robert J.; Salazar, Diego; Baer, Christina; Reinhardt, Jason; Priest, Galen; Barnett, Kirk (2016). "Ode to Ehrlich and Raven or how herbivorous insects might drive plant speciation". Ecology. 97 (11): 2939–2951. doi: 10.1002/ecy.1534 . ISSN   1939-9170. PMID   27870033 . Retrieved 2020-10-18.
  10. Thompson, John N. "Coevolution: the geographic mosaic of coevolutionary arms races." Current Biology 15.24 (2005): R992-R994.
  11. "Coevolution." Interactions Among Species. N.p., n.d. Web. 4 Nov. 1993. http://neko.bio.utk.edu/~ijuric/EEB460_%5B%5D
  12. Insect Ecology: Behavior, Populations and Communities By R. F. Denno, M. D. Eubanks
  13. Thompson, John N. "Coevolution." eLS (2001).
  14. Farrell, Brian D., David E. Dussourd, and Charles Mitter. "Escalation of plant defense: do latex and resin canals spur plant diversification?." American Naturalist (1991): 881-900.
  15. Kursar, Thomas A., and Phyllis D. Coley. "Convergence in defense syndromes of young leaves in tropical rainforests." Biochemical Systematics and Ecology 31.8 (2003): 929-949.
  16. Agrawal, Anurag A. "Natural selection on common milkweed (Asclepias syriaca) by a community of specialized insect herbivores." Evolutionary Ecology Research 7.5 (2005): 651.
  17. Engel, Katharina, and Ralph Tollrian. "Inducible defences as key adaptations for the successful invasion of Daphnia lumholtzi in North America?." Proceedings of the Royal Society B: Biological Sciences 276.1663 (2009): 1865-1873.
  18. Best, A., et al. "The Evolution of Host‐Parasite Range." The American naturalist 176.1 (2010): 63-71.