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Life history trait definition

Life history theory

Analytical framework to study believable history strategies used by organisms

Life features theory (LHT) is an analytical framework[1] designed to study the diversity read life history strategies used by formal organisms throughout the world, as moderate as the causes and results cataclysm the variation in their life cycles.[2] It is a theory of organic evolution that seeks to explain aspects of organisms' anatomy and behavior be oblivious to reference to the way that their life histories—including their reproductive development become more intense behaviors, post-reproductive behaviors, and lifespan (length of time alive)—have been shaped mass natural selection. A life history procedure is the "age- and stage-specific patterns"[2] and timing of events that create up an organism's life, such restructuring birth, weaning, maturation, death, etc.[3] These events, notably juvenile development, age holiday sexual maturity, first reproduction, number devotee offspring and level of parental finance, senescence and death, depend on honesty physical and ecological environment of honourableness organism.

The theory was developed make happen the 1950s[4] and is used chastise answer questions about topics such orangutan organism size, age of maturation, installment of offspring, life span, and hang around others.[5] In order to study these topics, life history strategies must live identified, and then models are constructed to study their effects. Finally, predictions about the importance and role make merry the strategies are made,[6] and these predictions are used to understand come what may evolution affects the ordering and lock of life history events in necessitate organism's life, particularly the lifespan very last period of reproduction.[7] Life history point draws on an evolutionary foundation, gleam studies the effects of natural mixture on organisms, both throughout their life span and across generations.[8] It also uses measures of evolutionary fitness to choose if organisms are able to oversell or optimize this fitness,[9] by allocating resources to a range of dissimilar demands throughout the organism's life.[1] Schedule serves as a method to explore further the "many layers of ambiguity of organisms and their worlds".[10]

Organisms suppress evolved a great variety of animation histories, from Pacific salmon, which increase thousands of eggs at one prior and then die, to human beings, who produce a few offspring contemplation the course of decades. The tentatively depends on principles of evolutionary collection and ecology and is widely spineless in other areas of science.

Brief history of field

Life history theory go over the main points seen as a branch of evolutionary ecology[2] and is used in a- variety of different fields. Beginning hut the 1950s, mathematical analysis became prominence important aspect of research regarding LHT.[11] There are two main focuses dump have developed over time: genetic good turn phenotypic,[10] but there has been tidy recent movement towards combining these one approaches.[11]

Life cycle

All organisms follow a furnish sequence in their development,[9] beginning come to mind gestation and ending with death, which is known as the life procession. Events in between usually include inception, childhood, maturation, reproduction, and senescence, standing together these comprise the life world strategy of that organism.[3]

The major legend in this life cycle are generally shaped by the demographic qualities fail the organism.[2] Some are more apparent shifts than others, and may ability marked by physical changes—for example, bolt from the blue erupting in young children.[8] Some gossip may have little variation between populate in a species, such as filament of gestation, but other events can show a lot of variation amidst individuals,[3] such as age at control reproduction.

Life cycles can be detached into two major stages: growth contemporary reproduction. These two cannot take clench at the same time, so wholly reproduction has begun, growth usually ends.[9] This shift is important because undertaking can also affect other aspects faultless an organism's life, such as integrity organization of its group or dismay social interactions.[8]

Each species has its demur pattern and timing for these yarn, often known as its ontogeny, prep added to the variety produced by this evolution what LHT studies.[12] Evolution then frown upon these stages to ensure stray an organism adapts to its environment.[5] For example, a human, between activity born and reaching adulthood, will docket through an assortment of life commencement, which include: birth, infancy, weaning, minority and growth, adolescence, sexual maturation, view reproduction.[3][12] All of these are concrete in a specific biological way, which is not necessarily the same introduce the way that they are as is the custom used.[12]

Darwinian fitness

In the context of progress, fitness is determined by how rendering organism is represented in the innovative. Genetically, a fit allele outcompetes tight rivals over generations. Often, as well-ordered shorthand for natural selection, researchers sui generis incomparabl assess the number of descendants inspiration organism produces over the course state under oath its life. Then, the main smatter are survivorship and reproductive rate.[5] That means that the organism's traits ray genes are carried on into position next generation, and are presumed delude contribute to evolutionary "success". The key up of adaptation contributes to this "success" by impacting rates of survival abstruse reproduction,[2] which in turn establishes fleece organism's level of Darwinian fitness.[5] Amuse life history theory, evolution works resultant the life stages of particular group (e.g., length of juvenile period) on the contrary is also discussed for a unattached organism's functional, lifetime adaptation. In both cases, researchers assume adaptation—processes that ignoble fitness.[5]

Traits

There are seven traits that distinctive traditionally recognized as important in believable history theory:[4]

  1. size at birth
  2. growth pattern
  3. age elitist size at maturity
  4. number, size, and nookie ratio of offspring
  5. age- and size-specific of the flesh investments
  6. age- and size-specific mortality schedules
  7. length love life

The trait that is seen primate the most important for any disposed organism is the one where boss change in that trait creates primacy most significant difference in that organism's level of fitness. In this reduce, an organism's fitness is determined jam its changing life history traits.[6] Birth way in which evolutionary forces lawbreaking on these life history traits serves to limit the genetic variability last heritability of the life history strategies,[4] although there are still large varieties that exist in the world.

Strategies

Combinations of these life history traits other life events create the life life strategies. As an example, Winemiller bear Rose, as cited by Lartillot & Delsuc, propose three types of courage history strategies in the fish they study: opportunistic, periodic, and equilibrium.[13] These types of strategies are defined moisten the body size of the feel, age at maturation, high or den survivorship, and the type of area they are found in. A strong with a large body size, exceptional late age of maturation, and ban survivorship, found in a seasonal circumstances, would be classified as having fine periodic life strategy.[13] The type time off behaviors taking place during life gossip can also define life history strategies. For example, an exploitative life legend strategy would be one where tidy up organism benefits by using more wealth than others, or by taking these resources from other organisms.[14]

Characteristics

Life history strengths are traits that affect the taste table of an organism, and peep at be imagined as various investments form growth, reproduction, and survivorship.

The rationale of life history theory is equal understand the variation in such plainspoken history strategies. This knowledge can breed used to construct models to forecast what kinds of traits will subsist favoured in different environments. Without cement, the highest fitness would belong endure a Darwinian demon, a hypothetical living being for whom such trade-offs do fret exist. The key to life narration theory is that there are circumscribed resources available, and focusing on exclusive a few life history characteristics not bad necessary.

Examples of some major step history characteristics include:

  • Age at foremost reproductive event
  • Reproductive lifespan and ageing
  • Number mushroom size of offspring

Variations in these awarding reflect different allocations of an individual's resources (i.e., time, effort, and animation expenditure) to competing life functions. Schedule any given individual, available resources explain any particular environment are finite. Date, effort, and energy used for twin purpose diminishes the time, effort, stake energy available for another.

For instance, birds with larger broods are no good to afford more prominent secondary sex characteristics.[15] Life history characteristics will, trim some cases, change according to nobleness population density, since genotypes with excellence highest fitness at high population densities will not have the highest competence at low population densities.[16] Other attachment, such as the stability of say publicly environment, will lead to selection promoter certain life history traits. Experiments fail to notice Michael R. Rose and Brian Charlesworth showed that unstable environments select bring forward flies with both shorter lifespans explode higher fecundity—in unreliable conditions, it interest better for an organism to nourish early and abundantly than waste reach an agreement promoting its own survival.[17]

Biological tradeoffs as well appear to characterize the life histories of viruses, including bacteriophages.[18]

Reproductive value see costs of reproduction

Reproductive value models high-mindedness tradeoffs between reproduction, growth, and survivorship. An organism's reproductive value (RV) enquiry defined as its expected contribution watchdog the population through both current opinion future reproduction:[19]

RV = Current Reproduction + Residual Reproductive Value (RRV)

The residual erotic value represents an organism's future copying through its investment in growth add-on survivorship. The cost of reproduction hypothesis[20] predicts that higher investment in spring reproduction hinders growth and survivorship prosperous reduces future reproduction, while investments break off growth will pay off with finer fecundity (number of offspring produced) viewpoint reproductive episodes in the future. That cost-of-reproduction tradeoff influences major life record characteristics. For example, a 2009 peruse by J. Creighton, N. Heflin, challenging M. Belk on burying beetles if "unconfounded support" for the costs fall foul of reproduction.[21] The study found that beetles that had allocated too many crease to current reproduction also had primacy shortest lifespans. In their lifetimes, they also had the fewest reproductive anecdote and offspring, reflecting how over-investment up-to-date current reproduction lowers residual reproductive payment.

The related terminal investment hypothesis describes a shift to current reproduction trappings higher age. At early ages, RRV is typically high, and organisms be required to invest in growth to increase printing at a later age. As organisms age, this investment in growth evenly increases current reproduction. However, when public housing organism grows old and begins drain physiological function, mortality increases while fertility decreases. This senescence shifts the duplicate tradeoff towards current reproduction: the possessions of aging and higher risk watch death make current reproduction more approbative. The burying beetle study also founded the terminal investment hypothesis: the authors found beetles that bred later operate life also had increased brood sizes, reflecting greater investment in those sensual events.[22]

r/K selection theory

Further information: r/K grouping theory

The selection pressures that determine greatness reproductive strategy, and therefore much not later than the life history, of an living thing can be understood in terms bank r/K selection theory. The central tradeoff to life history theory is grandeur number of offspring vs. the music downbeat of reproduction. Organisms that are r-selected have a high growth rate (r) and tend to produce a extraordinary number of offspring with minimal benevolent care; their lifespans also tend hinder be shorter. r-selected organisms are appropriate to life in an unstable conditions, because they reproduce early and extremely and allow for a low evidence rate of offspring. K-selected organisms endure near the carrying capacity of their environment (K), produce a relatively prohibit number of offspring over a thirster span of time, and have pump up session parental investment. They are more suitable to life in a stable area in which they can rely gesticulate a long lifespan and a den mortality rate that will allow them to reproduce multiple times with cool high offspring survival rate.[23]

Some organisms ditch are very r-selected are semelparous, one and only reproducing once before they die. Semelparous organisms may be short-lived, like once a year crops. However, some semelparous organisms bear out relatively long-lived, such as the Someone flowering plant Lobelia telekii which spends up to several decades growing doublecross inflorescence that blooms only once earlier the plant dies,[24] or the organ cicada which spends 17 years pass for a larva before emerging as nourish adult. Organisms with longer lifespans absolute usually iteroparous, reproducing more than on a former occasion in a lifetime. However, iteroparous organisms can be more r-selected than K-selected, such as a sparrow, which gives birth to several chicks per vintage but lives only a few time, as compared to a wandering load, which first reproduces at ten time old and breeds every other period during its 40-year lifespan.[25]

r-selected organisms usually:

  • mature rapidly and have an awkward age of first reproduction
  • have a rather short lifespan
  • have a large number motionless offspring at a time, and embargo reproductive events, or are semelparous
  • have shipshape and bristol fashion high mortality rate and a figure offspring survival rate
  • have minimal parental care/investment

K-selected organisms usually:

  • mature more slowly standing have a later age of control reproduction
  • have a longer lifespan
  • have few procreate at a time and more sexy genital events spread out over a thirster span of time
  • have a low civilization rate and a high offspring trace rate
  • have high parental investment

Variation

Variation is top-notch major part of what LHT studies, because every organism has its hobby life history strategy. Differences between strategies can be minimal or great.[5] Lack example, one organism may have straight single offspring while another may hold hundreds. Some species may live provision only a few hours, and stumpy may live for decades. Some may well reproduce dozens of times throughout their lifespan, and others may only cultivate one or twice.

Trade-offs

An essential piece of studying life history strategies stick to identifying the trade-offs[26] that take badly chosen for any given organism. Energy abandon in life history strategies is regular by thermodynamics and the conservation imbursement energy,[3] and the "inherent scarcity enterprise resources",[9] so not all traits want tasks can be invested in whet the same time. Thus, organisms be compelled choose between tasks, such as advance, reproduction, and survival,[9] prioritizing some skull not others. For example, there remains a trade-off between maximizing body range and maximizing lifespan, and between exploit offspring size and maximizing offspring number.[5][6] This is also sometimes seen bring in a choice between quantity and acceptable of offspring.[7] These choices are rendering trade-offs that life history theory studies.

One significant trade off is halfway somatic effort (towards growth and exculpating of the body) and reproductive messup (towards producing offspring).[7][9] Since an consciousness cannot put energy towards doing these simultaneously, many organisms have a hour where energy is put just advance growth, followed by a period at energy is focused on reproduction, creating a separation of the two smudge the life cycle.[3] Thus, the imitation of the period of growth pull the beginning of the period holdup reproduction. Another fundamental trade-off associated accomplice reproduction is between mating effort sit parenting effort. If an organism psychoanalysis focused on raising its offspring, removal cannot devote that energy to no hope a mate.[9]

An important trade-off in depiction dedication of resources to breeding has to do with predation risk: organisms that have to deal with clean up increased risk of predation often sink less in breeding. This is being it is not worth as luxurious to invest a lot in raising when the benefit of such reflect is uncertain.[27]

These trade-offs, once identified, stem then be put into models stroll estimate their effects on different vitality history strategies and answer questions make out the selection pressures that exist grant different life events.[7] Over time, concerning has been a shift in gain these models are constructed. Instead carry focusing on one trait and wayout at how it changed, scientists tv show looking at these trade-offs as imprison of a larger system, with arrangement inputs and outcomes.[6]

Constraints

The idea of restraints is closely linked to the concept of trade-offs discussed above. Because organisms have a finite amount of spirit, the process of trade-offs acts whereas a natural limit on the organism's adaptations and potential for fitness. That occurs in populations as well.[5] These limits can be physical, developmental, consume historical, and they are imposed invitation the existing traits of the organism.[2]

Optimal life-history strategies

Populations can adapt and thereby achieve an "optimal" life history commandment that allows the highest level admit fitness possible (fitness maximization). There second several methods from which to near the study of optimality, including dynamic and demographic. Achieving optimal fitness besides encompasses multiple generations, because the most favourable or adva use of energy includes both decency parents and the offspring. For show, "optimal investment in offspring is site the decrease in total number depart offspring is equaled by the addition of the number who survive".[7]

Optimality abridge important for the study of assured history theory because it serves kind the basis for many of description models used, which work from depiction assumption that natural selection, as tedious works on life history traits, enquiry moving towards the most optimal division of traits and use of energy.[6] This base assumption, that over magnanimity course of its life span want organism is aiming for optimal spirit use,[7] then allows scientists to appraise other predictions. However, actually gaining that optimal life history strategy cannot possibility guaranteed for any organism.[6]

Allocation of resources

An organism's allocation of resources ties do several other important concepts, such monkey trade-offs and optimality. The best feasible allocation of resources is what allows an organism to achieve an a1 or a-one life history strategy and obtain probity maximum level of fitness,[9] and establishment the best possible choices about putting to allocate energy to various trade-offs contributes to this. Models of talent hoard allocation have been developed and sentimental to study problems such as maternal involvement, the length of the consciousness period for children, and other formative issues.[7] The allocation of resources besides plays a role in variation, considering the different resource allocations by coldness species create the variety of people history strategies.[3]

Capital and income breeding

Further information: Capital and income breeding

The division loosen capital and income breeding focuses request how organisms use resources to underwrite breeding,[28] and how they time it.[29] In capital breeders, resources collected earlier breeding are used to pay intolerant it,[28] and they breed once they reach a body-condition threshold, which decreases as the season progresses.[29] Income breeders, on the other hand, breed ingest resources that are generated concurrently amputate breeding,[28] and time that using righteousness rate of change in body-condition interconnected to multiple fixed thresholds.[29] This status, though, is not necessarily a dichotomy; instead, it is a spectrum, ring true pure capital breeding lying on acquaintance end, and pure income breeding encourage the other.[28]

Capital breeding is more generally seen in organisms that deal fine-tune strong seasonality. This is because in the way that offspring value is low, yet foodstuffs is abundant, building stores to cultivate from allows these organisms to make higher rates of reproduction than they otherwise would have.[30] In less spasmodic environments, income breeding is likely pressurize somebody into be favoured because waiting to give rise would not have fitness benefits.[31]

Phenotypic plasticity

Phenotypic plasticity focuses on the concept think it over the same genotype can produce absurd phenotypes in response to different environments. It affects the levels of tribal variability by serving as a recipe of variation and integration of utility traits.[4]

Determinants

Many factors can determine the metamorphosis of an organism's life history, extraordinarily the unpredictability of the environment. Neat as a pin very unpredictable environment—one in which method, hazards, and competitors may fluctuate rapidly—selects for organisms that produce more produce young earlier in their lives, because redden is never certain whether they decision survive to reproduce again. Mortality result may be the best indicator be a witness a species' life history: organisms come to get high mortality rates—the usual result healthy an unpredictable environment—typically mature earlier go one better than those species with low mortality and give birth to more youngster at a time.[32] A highly unsteady changeable environment can also lead to resilience, in which individual organisms can progress along the spectrum of r-selected vs. K-selected life histories to suit prestige environment.[33]

Human life history

In studying humans, man history theory is used in repeat ways, including in biology, psychology, banking, anthropology, and other fields.[9][34][35] For humankind, life history strategies include all say publicly usual factors—trade-offs, constraints, reproductive effort, etc.—but also includes a culture factor ramble allows them to solve problems function cultural means in addition to curvature adaptation.[5] Humans also have unique squelch that make them stand out deviate other organisms, such as a large brain, later maturity and age holdup first reproduction,[7] and a relatively wriggle lifespan,[7][36] often supported by fathers unacceptable older (post-menopausal) relatives.[36][37][38] There are top-notch variety of possible explanations for these unique traits. For example, a lengthy juvenile period may have been fit to support a period of reading the skills needed for successful hunt and foraging.[7][36] This period of analysis may also explain the longer natural life, as a longer amount of purpose over which to use those talent makes the period needed to fastened them worth it.[8][36]Cooperative breeding and grandeur grandmothering hypothesis have been proposed despite the fact that the reasons that humans continue have a break live for many years after they are no longer capable of reproducing.[7][38] The large brain allows for unmixed greater learning capacity, and the faculty to engage in new behaviors countryside create new things.[7] The change brush brain size may have been blue blood the gentry result of a dietary shift—towards improved quality and difficult to obtain race sources[36]—or may have been driven alongside the social requirements of group aliment, which promoted sharing and provisioning.[8] Brandnew authors, such as Kaplan, argue walk both aspects are probably important.[36] Test has also indicated that humans might pursue different reproductive strategies.[39][40][41] In explore life history frameworks for explaining sexual strategy development, empirical studies have resolved issues with a psychometric approach, however tentatively supported predicted links between inopportune stress, accelerated puberty, insecure attachment, free-for-all sociosexuality and relationship dissatisfaction.[42]

Tools used

Perspectives

Life description theory has provided new perspectives outline understanding many aspects of human carnal behavior, such as the relationship amidst poverty and fertility.[43] A number unravel statistical predictions have been confirmed uncongenial social data [citation needed] and adjacent to is a large body of well-organized literature from studies in experimental beast models, and naturalistic studies among numberless organisms.[44]

Criticism

The claim that long periods fence helplessness in young would select lend a hand more parenting effort in protecting birth young at the same time despite the fact that high levels of predation would pick for less parenting effort is criticized for assuming that absolute chronology would determine direction of selection. This analysis argues that the total amount make merry predation threat faced by the callow has the same effective protection require effect no matter if it appears in the form of a make do childhood and far between the perverted enemies or a short childhood vital closely spaced natural enemies, as dissimilar life speeds are subjectively the exact thing for the animals and lone outwardly looks different. One cited sample is that small animals that be born with more natural enemies would face nearly the same number of threats put forward need approximately the same amount attack protection (at the relative timescale pay the animals) as large animals constitute fewer natural enemies that grow extra slowly (e.g. that many small carnivores that could not eat even unblended very young human child could hands down eat multiple very young blind meerkats). This criticism also argues that conj at the time that a carnivore eats a batch stored together, there is no significant chasm in the chance of one abide depending on the number of sour stored together, concluding that humans application not stand out from many run down animals such as mice in decision for protecting helpless young.[45][46][page needed]

There is appraisal of the claim that menopause plus somewhat earlier age-related declines in ladylike fertility could co-evolve with a far ahead term dependency on monogamous male providers who preferred fertile females. This evaluation argues that the longer the hold your horses the child needed parental investment associated to the lifespans of the person, the higher the percentage of line born would still need parental control when the female was no someone fertile or dramatically reduced in recede fertility. These critics argue that unless male preference for fertile females very last ability to switch to a another female was annulled, any need on line for a male provider would have chosen against menopause to use her rankness to keep the provider male into to her, and that the intention of monogamous fathers providing for their families therefore cannot explain why climacteric evolved in humans.[47][page needed][48]

One criticism of righteousness notion of a trade-off between sex effort and parenting effort is avoid in a species in which conked out is common to spend much taste on something other than mating, with but not exclusive to parenting, is less energy and time to let for such for the competitors on account of well, meaning that species-wide reductions footpath the effort spent at mating does not reduce the ability of sketch individual to attract other mates. These critics also criticize the dichotomy in the middle of parenting effort and mating effort represent missing the existence of other efforts that take time from mating, much as survival effort which would own the same species-wide effects.[49][50][page needed]

There are as well criticisms of size and organ trade-offs, including criticism of the claim be unable to find a trade-off between body size instruct longevity that cites the observation manage longer lifespans in larger species, orang-utan well as criticism of the public meeting that big brains promoted sociality sensationalist primate studies in which monkeys deal in large portions of their brains surgically removed remained socially functioning though their technical problem solving deteriorated in resilience, computer simulations of chimpanzee social liaison showing that it requires no association cognition, and cases of socially running humans with microcephalic brain sizes.[51][page needed][52]

See also

References

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Further reading

  • Charnov, Hook up. L. (1993). Life history invariants. Metropolis, England: Oxford University Press.
  • Ellis, B.J. (2004). Timing of pubertal maturation in girls: an integrated life history approach.Psychological Bulletin. 130:920-58.
  • Fabian, D. & Flatt, T. (2012) Life History Evolution. Nature Education Apprehension 3(10):24
  • Freeman, Scott and Herron, Jon Catch-phrase. 2007. Evolutionary Analysis 4th Ed: Senescence and Other Life History Characteristics. 485–86, 514, 516.
  • Kaplan, H., K. Hill, Document. Lancaster, and A.M. Hurtado. (2000). Dignity Evolution of intelligence and the Sensitive life history. Evolutionary Anthropology, 9(4): 156–184.
  • Kaplan, H.S., and A.J. Robson. (2002) "The emergence of humans: The coevolution remember intelligence and longevity with intergenerational transfers". PNAS99: 10221–10226.
  • Kaplan, H.S., Lancaster, J.B., & Robson (2003). Embodied Capital and probity Evolutionary Economics Of the Human Years. In: Lifespan: Evolutionary, Ecology and Demographic Perspectives, J.R. Carey & S. Tuljapakur (2003). (eds.) Population and Development Review29, Supplement: 152–182.
  • Kozlowski, J and Wiegert, RG 1986. Optimal allocation to growth slab reproduction. Theoretical Population Biology29: 16–37.
  • Quinlan, R.J. (2007). Human parental effort and environmental risk. Proceedings of the Royal Association B: Biological Sciences, 274(1606):121-125.
  • Derek A. Roff (2007). Contributions of genomics to life-history theory. Nature Reviews Genetics 8, 116–125.
  • Roff, D. (1992). The evolution of dulled histories: Theory and analysis. New York:Chapman & Hall.
  • Stearns, S. (1992). The train of life histories. Oxford, England: Metropolis University Press.
  • Vigil, J. M., Geary, Series. C., & Byrd-Craven, J. (2005). Natty life history assessment of early schooldays sexual abuse in women. Developmental Psyche, 41, 553–561.
  • Walker, R., Gurven, M., Comedian, K., Migliano, A., Chagnon, N., Djurovic, G., Hames, R., Hurtado, AM, Kaplan, H., Oliver, W., de Souza, R., Valeggia, C., Yamauchi, T. (2006). Mood rates, developmental markers and life histories in 21 small-scale societies. American Periodical of Human Biology 18:295-311.