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161-402: Advocates of construction grammar argue that language and culture are not designed by people, but are 'emergent' or automatically constructed in a process which is comparable to natural selection in species or the formation of natural constructions such as nests made by social insects . Constructions correspond to replicators or memes in memetics and other cultural replicator theories. It

322-420: A synonymous substitution . However, many mutations in non-coding DNA have deleterious effects. Although both mutation rates and average fitness effects of mutations are dependent on the organism, a majority of mutations in humans are slightly deleterious. Some mutations occur in "toolkit" or regulatory genes . Changes in these often have large effects on the phenotype of the individual because they regulate

483-457: A "Nature" which would do the selection. At the time, other mechanisms of evolution such as evolution by genetic drift were not yet explicitly formulated, and Darwin believed that selection was likely only part of the story: "I am convinced that Natural Selection has been the main but not exclusive means of modification." In a letter to Charles Lyell in September 1860, Darwin regretted the use of

644-702: A 'recent' selective sweep near the centre of the block. Background selection is the opposite of a selective sweep. If a specific site experiences strong and persistent purifying selection, linked variation tends to be weeded out along with it, producing a region in the genome of low overall variability. Because background selection is a result of deleterious new mutations, which can occur randomly in any haplotype, it does not produce clear blocks of linkage disequilibrium, although with low recombination it can still lead to slightly negative linkage disequilibrium overall. Prosody (linguistics) In linguistics , prosody ( / ˈ p r ɒ s ə d i , ˈ p r ɒ z -/ )

805-519: A chromosome. During the formation of gametes, recombination reshuffles the alleles. The chance that such a reshuffle occurs between two alleles is inversely related to the distance between them. Selective sweeps occur when an allele becomes more common in a population as a result of positive selection. As the prevalence of one allele increases, closely linked alleles can also become more common by " genetic hitchhiking ", whether they are neutral or even slightly deleterious. A strong selective sweep results in

966-847: A computational model for large scale practical usage in Natural Language Processing frameworks but interest in construction grammar has been shown by more traditional computational linguists as a contrast to the current boom in more opaque deep learning models. This is largely due to the representational convenience of CxG models and their potential to integrate with current tokenizers as a perceptual layer for further processing in neurally inspired models. Approaches to integrate constructional grammar with existing Natural Language Processing frameworks include hand-built feature sets and templates and used computational models to identify their prevalence in text collections, but some suggestions for more emergent models have been proposed, e.g. in

1127-554: A different, incompatible allele of the same gene. Selection against the heterozygote would then directly create reproductive isolation, leading to the Bateson–Dobzhansky–Muller model , further elaborated by H. Allen Orr and Sergey Gavrilets . With reinforcement , however, natural selection can favor an increase in pre-zygotic isolation, influencing the process of speciation directly. Natural selection acts on an organism's phenotype, or physical characteristics. Phenotype

1288-503: A group of lexemes. Combinatorial constructions include both inflectional and derivational constructions. SBCG is both formal and generative; while cognitive-functional grammarians have often opposed their standards and practices to those of formal, generative grammarians, there is in fact no incompatibility between a formal, generative approach and a rich, broad-coverage, functionally based grammar. It simply happens that many formal, generative theories are descriptively inadequate grammars. SBCG

1449-435: A learned association of form and prototypical meaning in order to set up the constructions said to form the basic units of syntax. Natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype . It is a key mechanism of evolution , the change in the heritable traits characteristic of a population over generations. Charles Darwin popularised

1610-549: A model merely of idioms or infrequently used, minor patterns. As the pairing of the syntactic construction and its prototypical meaning are learned in early childhood, children should initially learn the basic constructions with their prototypical semantics, that is, 'agent of action' for the subject in the SV relation, 'affected object of agent's action' for the direct object term in VO, and 'recipient in transfer of possession of object' for

1771-462: A neutralization of semantic distinctions. For instance, in a detailed discussion of the dissociation of grammatical case-roles from semantics, Talmy Givon lists the multiple semantic roles of subjects and direct objects in English. As these phenomena are well-established, some linguists propose that core grammatical relations be excluded from CxG as they are not constructions, leaving the theory to be

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1932-685: A population if the environment that favours these traits remains fixed. If new traits become more favored due to changes in a specific niche , microevolution occurs. If new traits become more favored due to changes in the broader environment, macroevolution occurs. Sometimes, new species can arise especially if these new traits are radically different from the traits possessed by their predecessors. The likelihood of these traits being 'selected' and passed down are determined by many factors. Some are likely to be passed down because they adapt well to their environments. Others are passed down because these traits are actively preferred by mating partners, which

2093-479: A precursor to speciation . Alternatively, selection can be divided according to its effect on genetic diversity . Purifying or negative selection acts to remove genetic variation from the population (and is opposed by de novo mutation , which introduces new variation. In contrast, balancing selection acts to maintain genetic variation in a population, even in the absence of de novo mutation, by negative frequency-dependent selection . One mechanism for this

2254-402: A question. In the second line, pitch falls, indicating a statement ‍ — a confirmation of the first line in this case. Finally, in the third line, a complicated rise-fall pattern indicates incredulity. Each pitch/intonation pattern communicates a different meaning. An additional pitch-related variation is pitch range ; speakers are capable of speaking with a wide range of pitch (this

2415-441: A region of the genome where the positively selected haplotype (the allele and its neighbours) are in essence the only ones that exist in the population. Selective sweeps can be detected by measuring linkage disequilibrium , or whether a given haplotype is overrepresented in the population. Since a selective sweep also results in selection of neighbouring alleles, the presence of a block of strong linkage disequilibrium might indicate

2576-430: A relatively high probability of surviving to adulthood. Natural selection can act on any heritable phenotypic trait , and selective pressure can be produced by any aspect of the environment, including sexual selection and competition with members of the same or other species. However, this does not imply that natural selection is always directional and results in adaptive evolution; natural selection often results in

2737-458: A reproductive advantage regardless of whether or not the trait is heritable. Following Darwin's primary usage, the term is used to refer both to the evolutionary consequence of blind selection and to its mechanisms. It is sometimes helpful to explicitly distinguish between selection's mechanisms and its effects; when this distinction is important, scientists define "(phenotypic) natural selection" specifically as "those mechanisms that contribute to

2898-482: A speaker is inviting the listener to make a contribution to the conversation. Prosody is also important in signalling emotions and attitudes. When this is involuntary (as when the voice is affected by anxiety or fear), the prosodic information is not linguistically significant. However, when the speaker varies their speech intentionally, for example to indicate sarcasm, this usually involves the use of prosodic features. The most useful prosodic feature in detecting sarcasm

3059-454: A species were uninteresting departures from their Platonic ideals (or typus ) of created kinds . However, the theory of uniformitarianism in geology promoted the idea that simple, weak forces could act continuously over long periods of time to produce radical changes in the Earth's landscape. The success of this theory raised awareness of the vast scale of geological time and made plausible

3220-438: A uniform mechanism for parsing and production. Moreover, it has been demonstrated through robotic experiments that FCG grammars can be grounded in embodiment and sensorimotor experiences. FCG integrates many notions from contemporary computational linguistics such as feature structures and unification-based language processing. Constructions are considered bidirectional and hence usable both for parsing and production. Processing

3381-536: A variety of "filled" pause types. Formulaic language pause fillers include "Like", "Er" and "Um", and paralinguistic expressive respiratory pauses include the sigh and gasp . Although related to breathing, pauses may contain contrastive linguistic content, as in the periods between individual words in English advertising voice-over copy sometimes placed to denote high information content, e.g. "Quality. Service. Value". Pausing or its lack contributes to

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3542-483: A well-documented example involves the spread of a gene in the butterfly Hypolimnas bolina suppressing male-killing activity by Wolbachia bacteria parasites on the island of Samoa , where the spread of the gene is known to have occurred over a period of just five years. A prerequisite for natural selection to result in adaptive evolution, novel traits and speciation is the presence of heritable genetic variation that results in fitness differences. Genetic variation

3703-438: A word. Take one popular English word for example: In English, lexical prosody is used for a few different reasons. As we have seen above, lexical prosody was used to change the form of a word from a noun to a verb. Another function of lexical prosody has to do with the grammatical role that a word plays within a sentence. Adjectives and nouns of a sentence are often stressed on the first syllables while verbs are often stressed on

3864-532: Is heterozygote advantage , where individuals with two different alleles have a selective advantage over individuals with just one allele. The polymorphism at the human ABO blood group locus has been explained in this way. Another option is to classify selection by the life cycle stage at which it acts. Some biologists recognise just two types: viability (or survival) selection , which acts to increase an organism's probability of survival, and fecundity (or fertility or reproductive) selection, which acts to increase

4025-498: Is a dispute between the advocates of construction grammar and memetics , an evolutionary approach which adheres to the Darwinian view of language and culture. Advocates of construction grammar argue that memetics takes the perspective of intelligent design to cultural evolution while construction grammar rejects human free will in language construction; but, according to memetician Susan Blackmore , this makes construction grammar

4186-674: Is a reduction in the mean fundamental frequency relative to other speech for humor, neutrality, or sincerity. While prosodic cues are important in indicating sarcasm, context clues and shared knowledge are also important. Emotional prosody was considered by Charles Darwin in The Descent of Man to predate the evolution of human language : "Even monkeys express strong feelings in different tones – anger and impatience by low, – fear and pain by high notes." Native speakers listening to actors reading emotionally neutral text while projecting emotions correctly recognized happiness 62% of

4347-405: Is all the parts of one whole) happened like as if they were made for the sake of something, these were preserved, having been appropriately constituted by an internal spontaneity, and whatsoever things were not thus constituted, perished, and still perish. But Aristotle rejected this possibility in the next paragraph, making clear that he is talking about the development of animals as embryos with

4508-405: Is argued that children pay close attention to each utterance they hear, and gradually make generalizations based on the utterances they have heard. Because constructions are learned, they are expected to vary considerably across different languages. In construction grammar, as in general semiotics , the grammatical construction is a pairing of form and content. The formal aspect of a construction

4669-588: Is argued that construction grammar is not an original model of cultural evolution, but for essential part the same as memetics . Construction grammar is associated with concepts from cognitive linguistics that aim to show in various ways how human rational and creative behaviour is automatic and not planned. Construction grammar was first developed in the 1980s by linguists such as Charles Fillmore , Paul Kay , and George Lakoff , in order to analyze idioms and fixed expressions. Lakoff's 1977 paper "Linguistic Gestalts" put forward an early version of CxG, arguing that

4830-500: Is central to natural selection. In broad terms, individuals that are more "fit" have better potential for survival, as in the well-known phrase " survival of the fittest ", but the precise meaning of the term is much more subtle. Modern evolutionary theory defines fitness not by how long an organism lives, but by how successful it is at reproducing. If an organism lives half as long as others of its species, but has twice as many offspring surviving to adulthood, its genes become more common in

4991-675: Is described by a lexeme or construction. Recent SBCG works have expanded on the lexicalist model of idiomatically combining expressions sketched out in Sag 2012. The type of construction grammar associated with linguists like Goldberg and Lakoff looks mainly at the external relations of constructions and the structure of constructional networks. In terms of form and function, this type of construction grammar puts psychological plausibility as its highest desideratum. It emphasizes experimental results and parallels with general cognitive psychology. It also draws on certain principles of cognitive linguistics. In

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5152-474: Is described to be a natural component of language. The defining features of prosody that display the nuanced emotions of an individual differ across languages and cultures. Some writers (e.g., O'Connor and Arnold) have described intonation entirely in terms of pitch, while others (e.g., Crystal) propose that "intonation" is a combination of several prosodic variables. English intonation is often said to be based on three aspects: The choice of pitch movement and

5313-455: Is determined by an organism's genetic make-up (genotype) and the environment in which the organism lives. When different organisms in a population possess different versions of a gene for a certain trait, each of these versions is known as an allele . It is this genetic variation that underlies differences in phenotype. An example is the ABO blood type antigens in humans, where three alleles govern

5474-544: Is flexible in the sense that it can even cope with partially ungrammatical or incomplete sentences. FCG is called 'fluid' because it acknowledges the premise that language users constantly change and update their grammars. The research on FCG is conducted at Sony CSL Paris and the AI Lab at the Vrije Universiteit Brussel . Most of the above approaches to construction grammar have not been implemented as

5635-468: Is generative in a way that prevailing syntax-centered theories are not: its mechanisms are intended to represent all of the patterns of a given language, including idiomatic ones; there is no 'core' grammar in SBCG. SBCG a licensing-based theory, as opposed to one that freely generates syntactic combinations and uses general principles to bar illicit ones: a word, lexeme or phrase is well formed if and only if it

5796-540: Is important to distinguish between the personal characteristics that belong to an individual's voice (for example, their habitual pitch range, intonation patterns, etc.) and the independently variable prosodic features that are used contrastively to communicate meaning (for example, the use of changes in pitch to indicate the difference between statements and questions). Personal characteristics that belong to an individual are not linguistically significant while prosodic features are. Prosody has been found across all languages and

5957-456: Is increased, which means that it leaves more offspring. If the traits that give these individuals a reproductive advantage are also heritable , that is, passed from parent to offspring, then there will be differential reproduction, that is, a slightly higher proportion of fast rabbits or efficient algae in the next generation. Even if the reproductive advantage is very slight, over many generations any advantageous heritable trait becomes dominant in

6118-438: Is intrinsic to the concept of a species that hybrids are selected against, opposing the evolution of reproductive isolation, a problem that was recognised by Darwin. The problem does not occur in allopatric speciation with geographically separated populations, which can diverge with different sets of mutations. E. B. Poulton realized in 1903 that reproductive isolation could evolve through divergence, if each lineage acquired

6279-543: Is known as sexual selection . Female bodies also prefer traits that confer the lowest cost to their reproductive health, which is known as fecundity selection . Natural selection is a cornerstone of modern biology . The concept, published by Darwin and Alfred Russel Wallace in a joint presentation of papers in 1858 , was elaborated in Darwin's influential 1859 book On the Origin of Species by Means of Natural Selection, or

6440-483: Is lowered by the presence of another. This may be because both rely on a limited supply of a resource such as food, water, or territory . Competition may be within or between species , and may be direct or indirect. Species less suited to compete should in theory either adapt or die out , since competition plays a powerful role in natural selection, but according to the "room to roam" theory it may be less important than expansion among larger clades . Competition

6601-555: Is modelled by r/K selection theory , which is based on Robert MacArthur and E. O. Wilson 's work on island biogeography . In this theory, selective pressures drive evolution in one of two stereotyped directions: r - or K -selection. These terms, r and K , can be illustrated in a logistic model of population dynamics : d N d t = r N ( 1 − N K ) {\displaystyle {\frac {dN}{dt}}=rN\left(1-{\frac {N}{K}}\right)\qquad \!} where r

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6762-461: Is natural selection via any means other than sexual selection, such as kin selection, competition, and infanticide . Following Darwin, natural selection is sometimes defined as ecological selection, in which case sexual selection is considered a separate mechanism. Sexual selection as first articulated by Darwin (using the example of the peacock 's tail) refers specifically to competition for mates, which can be intrasexual , between individuals of

6923-685: Is necessary for listeners to be able to identify the affective tone of the utterance. At lengths below this, there was not enough information for listeners to process the emotional context of the utterance. Unique prosodic features have been noted in infant-directed speech (IDS) - also known as baby talk , child-directed speech (CDS), or "motherese". Adults, especially caregivers, speaking to young children tend to imitate childlike speech by using higher and more variable pitch, as well as an exaggerated stress. These prosodic characteristics are thought to assist children in acquiring phonemes, segmenting words, and recognizing phrasal boundaries. And though there

7084-426: Is no evidence to indicate that infant-directed speech is necessary for language acquisition, these specific prosodic features have been observed in many different languages. An aprosodia is an acquired or developmental impairment in comprehending or generating the emotion conveyed in spoken language. Aprosody is often accompanied by the inability to properly utilize variations in speech, particularly with deficits in

7245-812: Is often recognized, and that they are best understood as multi-word, partially filled constructions. Construction grammar rejects the idea that there is a sharp dichotomy between lexical items , which are arbitrary and specific, and grammatical rules, which are completely general. Instead, CxG posits that there are linguistic patterns at every level of generality and specificity: from individual words, to partially filled constructions (e.g. drive X crazy ), to fully abstract rules (e.g. subject–auxiliary inversion ). All of these patterns are recognized as constructions. In contrast to theories that posit an innate universal grammar for all languages, construction grammar holds that speakers learn constructions inductively as they are exposed to them, using general cognitive processes. It

7406-410: Is selection favoring extreme trait values over intermediate trait values. Disruptive selection may cause sympatric speciation through niche partitioning . Some forms of balancing selection do not result in fixation, but maintain an allele at intermediate frequencies in a population. This can occur in diploid species (with pairs of chromosomes) when heterozygous individuals (with just one copy of

7567-412: Is slowed by the competition. The variant which is a candidate for a beneficial mutation in this limited carrying capacity environment must first out-compete the "less fit" variants in order to accumulate the requisite number of replications for there to be a reasonable probability of that beneficial mutation occurring. In biology, competition is an interaction between organisms in which the fitness of one

7728-427: Is sometimes referred to as the accentual function of prosody. A well-known example is the ambiguous sentence "I never said she stole my money", where there are seven meaning changes depending on which of the seven words is vocally highlighted. Prosody helps convey many other pragmatic functions, including expressing attitudes (approval, uncertainty, dissatisfaction, and so on), flagging turn-taking intentions (to hold

7889-461: Is still often used by non-biologists, modern biologists avoid it because it is tautological if "fittest" is read to mean "functionally superior" and is applied to individuals rather than considered as an averaged quantity over populations. Natural selection relies crucially on the idea of heredity, but developed before the basic concepts of genetics . Although the Moravian monk Gregor Mendel ,

8050-563: Is stored in the taxonomies: Because construction grammar does not operate with surface derivations from underlying structures, it adheres to functionalist linguist Dwight Bolinger 's principle of no synonymy , on which Adele Goldberg elaborates in her book. This means that construction grammarians argue, for instance, that active and passive versions of the same proposition are not derived from an underlying structure, but are instances of two different constructions. As constructions are pairings of form and meaning, active and passive versions of

8211-413: Is that conceptual semantics is primary to the degree that form mirrors, or is motivated by, content. Langacker argues that even abstract grammatical units like part-of-speech classes are semantically motivated and involve certain conceptualizations. William A. Croft 's radical construction grammar is designed for typological purposes and takes into account cross-linguistic factors. It deals mainly with

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8372-499: Is that the content of all linguistic signs involves mental simulations and is ultimately dependent on basic image schemas of the kind advocated by Mark Johnson and George Lakoff, and so ECG aligns itself with cognitive linguistics. Like construction grammar, embodied construction grammar makes use of a unification-based model of representation. A non-technical introduction to the NTL theory behind embodied construction grammar as well as

8533-405: Is the growth rate of the population ( N ), and K is the carrying capacity of its local environmental setting. Typically, r -selected species exploit empty niches , and produce many offspring, each with a relatively low probability of surviving to adulthood. In contrast, K -selected species are strong competitors in crowded niches, and invest more heavily in much fewer offspring, each with

8694-405: Is the basis of the metric pattern, we have poetry; when pitch is the pattern basis, we have rhythmic prose" (Weeks 11). Stress retraction is a popular example of phrasal prosody in everyday life. For example: Contrastive stress is another everyday English example of phrasal prosody that helps us determine what parts of the sentence are important. Take these sentences for example: Emphasizing that

8855-483: Is the major factor, the resulting prominence is often called accent rather than stress. There is considerable variation from language to language concerning the role of stress in identifying words or in interpreting grammar and syntax. Although rhythm is not a prosodic variable in the way that pitch or loudness are, it is usual to treat a language's characteristic rhythm as a part of its prosodic phonology. It has often been asserted that languages exhibit regularity in

9016-435: Is the result of mutations, genetic recombinations and alterations in the karyotype (the number, shape, size and internal arrangement of the chromosomes ). Any of these changes might have an effect that is highly advantageous or highly disadvantageous, but large effects are rare. In the past, most changes in the genetic material were considered neutral or close to neutral because they occurred in noncoding DNA or resulted in

9177-462: Is the sign, with subtypes word, lexeme and phrase. The inclusion of phrase within the canon of signs marks a major departure from traditional syntactic thinking. In SBCG, phrasal signs are licensed by correspondence to the mother of some licit construct of the grammar. A construct is a local tree with signs at its nodes. Combinatorial constructions define classes of constructs. Lexical class constructions describe combinatoric and other properties common to

9338-428: Is the study of elements of speech that are not individual phonetic segments (vowels and consonants) but which are properties of syllables and larger units of speech, including linguistic functions such as intonation , stress , and rhythm . Such elements are known as suprasegmentals . Prosody reflects the nuanced emotional features of the speaker or of their utterances: their obvious or underlying emotional state,

9499-419: Is typically associated with the following: Some of these cues are more powerful or prominent than others. Alan Cruttenden, for example, writes "Perceptual experiments have clearly shown that, in English at any rate, the three features (pitch, length and loudness) form a scale of importance in bringing syllables into prominence, pitch being the most efficacious, and loudness the least so". When pitch prominence

9660-543: Is typically described as a syntactic template, but the form covers more than just syntax, as it also involves phonological aspects, such as prosody and intonation . The content covers semantic as well as pragmatic meaning. The semantic meaning of a grammatical construction is made up of conceptual structures postulated in cognitive semantics : image-schemas , frames, conceptual metaphors, conceptual metonymies, prototypes of various kinds, mental spaces, and bindings across these (called "blends"). Pragmatics just becomes

9821-454: Is used by listeners to guide decisions about the emotional affect of the situation. Whether a person decodes the prosody as positive, negative, or neutral plays a role in the way a person decodes a facial expression accompanying an utterance. As the facial expression becomes closer to neutral, the prosodic interpretation influences the interpretation of the facial expression. A study by Marc D. Pell revealed that 600 ms of prosodic information

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9982-616: Is usually associated with excitement), while at other times with a narrow range. English makes use of changes in key ; shifting one's intonation into the higher or lower part of one's pitch range is believed to be meaningful in certain contexts. Stress functions as the means of making a syllable prominent. Stress may be studied in relation to individual words (named "word stress" or lexical stress ) or in relation to larger units of speech (traditionally referred to as "sentence stress" but more appropriately named " prosodic stress "). Stressed syllables are made prominent by several variables. Stress

10143-474: The Industrial Revolution , many of the trees on which the moths rested became blackened by soot , giving the dark-coloured moths an advantage in hiding from predators. This gave dark-coloured moths a better chance of surviving to produce dark-coloured offspring, and in just fifty years from the first dark moth being caught, nearly all of the moths in industrial Manchester were dark. The balance

10304-551: The Roman poet Lucretius , expressed the idea that nature produces a huge variety of creatures, randomly, and that only those creatures that manage to provide for themselves and reproduce successfully persist. Empedocles' idea that organisms arose entirely by the incidental workings of causes such as heat and cold was criticised by Aristotle in Book II of Physics . He posited natural teleology in its place, and believed that form

10465-539: The ditransitive schema [S V IO DO] is said to express semantic content X CAUSES Y TO RECEIVE Z, just like kill means X CAUSES Y TO DIE. In construction grammar, a grammatical construction, regardless of its formal or semantic complexity and make up, is a pairing of form and meaning. Thus words and word classes may be regarded as instances of constructions. Indeed, construction grammarians argue that all pairings of form and meaning are constructions, including phrase structures, idioms , words and even morphemes . Unlike

10626-524: The founder effect in initially small new populations. When genetic variation does not result in differences in fitness, selection cannot directly affect the frequency of such variation. As a result, the genetic variation at those sites is higher than at sites where variation does influence fitness. However, after a period with no new mutations, the genetic variation at these sites is eliminated due to genetic drift. Natural selection reduces genetic variation by eliminating maladapted individuals, and consequently

10787-434: The genetic regulatory programs which control the development of the embryo at molecular level. Natural selection is here understood to act on embryonic development to change the morphology of the adult body. The term natural selection is most often defined to operate on heritable traits, because these directly participate in evolution. However, natural selection is "blind" in the sense that changes in phenotype can give

10948-436: The isochrony article, this claim has not been supported by scientific evidence. Voiced or unvoiced, the pause is a form of interruption to articulatory continuity such as an open or terminal juncture . Conversation analysis commonly notes pause length. Distinguishing auditory hesitation from silent pauses is one challenge. Contrasting junctures within and without word chunks can aid in identifying pauses. There are

11109-418: The left hemisphere, which contains Wernicke's area ). Damage to the right inferior frontal gyrus causes a diminished ability to convey emotion or emphasis by voice or gesture, and damage to right superior temporal gyrus causes problems comprehending emotion or emphasis in the voice or gestures of others. The right Brodmann area 22 aids in the interpretation of prosody, and damage causes sensory aprosodia, with

11270-424: The modern synthesis of the mid-20th century. The addition of molecular genetics has led to evolutionary developmental biology , which explains evolution at the molecular level. While genotypes can slowly change by random genetic drift , natural selection remains the primary explanation for adaptive evolution . Several philosophers of the classical era , including Empedocles and his intellectual successor,

11431-468: The "principle by which each slight variation [of a trait], if useful, is preserved". The concept was simple but powerful: individuals best adapted to their environments are more likely to survive and reproduce. As long as there is some variation between them and that variation is heritable , there will be an inevitable selection of individuals with the most advantageous variations. If the variations are heritable, then differential reproductive success leads to

11592-496: The "upper power" but instead are generated in different forms naturally and then selected for reproduction by their compatibility with the environment. The more recent classical arguments were reintroduced in the 18th century by Pierre Louis Maupertuis and others, including Darwin's grandfather, Erasmus Darwin . Until the early 19th century, the prevailing view in Western societies was that differences between individuals of

11753-500: The 2023 Georgetown University Roundtable on Linguistics. Esa Itkonen , who defends humanistic linguistics and opposes Darwinian linguistics, questions the originality of the work of Adele Goldberg, Michael Tomasello, Gilles Fauconnier , William Croft and George Lakoff. According to Itkonen, construction grammarians have appropriated old ideas in linguistics adding some false claims. For example, construction type and conceptual blending correspond to analogy and blend , respectively, in

11914-459: The Goldbergian strand, constructions interact with each other in a network via four inheritance relations: polysemy link, subpart link, metaphorical extension, and finally instance link. Sometimes, Ronald Langacker's cognitive grammar framework is described as a type of construction grammar. Cognitive grammar deals mainly with the semantic content of constructions, and its central argument

12075-433: The Origin of Species (1942). W. D. Hamilton conceived of kin selection in 1964. This synthesis cemented natural selection as the foundation of evolutionary theory, where it remains today. A second synthesis was brought about at the end of the 20th century by advances in molecular genetics , creating the field of evolutionary developmental biology ("evo-devo"), which seeks to explain the evolution of form in terms of

12236-678: The Preservation of Favoured Races in the Struggle for Life . He described natural selection as analogous to artificial selection, a process by which animals and plants with traits considered desirable by human breeders are systematically favoured for reproduction. The concept of natural selection originally developed in the absence of a valid theory of heredity; at the time of Darwin's writing, science had yet to develop modern theories of genetics. The union of traditional Darwinian evolution with subsequent discoveries in classical genetics formed

12397-566: The STAIRS is how the man went up. Emphasizing that it was a MAN who went up the stairs. It's important to note that the right hemisphere of the brain dominates one's perception of prosody. In contrast to left hemisphere damage where patterns of aphasias are present, patterns of aprosodias are present with damage to the left hemisphere. In patients with right hemisphere lesions, they are characterized as monotonous and as lacking variety in their tone and expression. They're also seen to struggle with

12558-533: The VP construction but contain no phrasal information in themselves. In the mid-2000s, several of the developers of BCG, including Charles Fillmore, Paul Kay, Ivan Sag and Laura Michaelis, collaborated in an effort to improve the formal rigor of BCG and clarify its representational conventions. The result was Sign Based Construction Grammar (SBCG). SBCG is based on a multiple-inheritance hierarchy of typed feature structures. The most important type of feature structure in SBCG

12719-448: The ability to accurately modulate pitch, loudness, intonation, and rhythm of word formation. This is seen sometimes in autistic individuals. The three main types of aprosodia are: Lexical prosody refers to the specific amplitudes, pitches, or lengths of vowels that are applied to specific syllables in words based on what the speaker wants to emphasize. The different stressors placed on individual syllables can change entire meanings of

12880-441: The adult population of the next generation. Though natural selection acts on individuals, the effects of chance mean that fitness can only really be defined "on average" for the individuals within a population. The fitness of a particular genotype corresponds to the average effect on all individuals with that genotype. A distinction must be made between the concept of "survival of the fittest" and "improvement in fitness". "Survival of

13041-551: The allele is fixed and the entire population shares the fitter phenotype. Far more common is stabilizing selection, which lowers the frequency of alleles that have a deleterious effect on the phenotype—that is, produce organisms of lower fitness. This process can continue until the allele is eliminated from the population. Stabilizing selection conserves functional genetic features, such as protein-coding genes or regulatory sequences , over time by selective pressure against deleterious variants. Disruptive (or diversifying) selection

13202-427: The allele) have a higher fitness than homozygous individuals (with two copies). This is called heterozygote advantage or over-dominance, of which the best-known example is the resistance to malaria in humans heterozygous for sickle-cell anaemia . Maintenance of allelic variation can also occur through disruptive or diversifying selection , which favours genotypes that depart from the average in either direction (that is,

13363-544: The antlers of stags , which are used in combat with other stags. More generally, intrasexual selection is often associated with sexual dimorphism , including differences in body size between males and females of a species. Natural selection is seen in action in the development of antibiotic resistance in microorganisms . Since the discovery of penicillin in 1928, antibiotics have been used to fight bacterial diseases. The widespread misuse of antibiotics has selected for microbial resistance to antibiotics in clinical use, to

13524-404: The apparently vestigial structure may retain a limited functionality, or may be co-opted for other advantageous traits in a phenomenon known as preadaptation . A famous example of a vestigial structure, the eye of the blind mole-rat , is believed to retain function in photoperiod perception. Speciation requires a degree of reproductive isolation —that is, a reduction in gene flow. However, it

13685-688: The articulation of adjacent word syllables, thereby changing the potential open junctures between words into closed junctures. Prosody has had a number of perceptually significant functions in English and other languages, contributing to the recognition and comprehension of speech. It is believed that prosody assists listeners in parsing continuous speech and in the recognition of words, providing cues to syntactic structure, grammatical boundaries and sentence type. Boundaries between intonation units are often associated with grammatical or syntactic boundaries; these are marked by such prosodic features as pauses and slowing of tempo, as well as "pitch reset" where

13846-442: The assumption that groups replicate and mutate in an analogous way to genes and individuals. There is an ongoing debate over the degree to which group selection occurs in nature. Finally, selection can be classified according to the resource being competed for. Sexual selection results from competition for mates. Sexual selection typically proceeds via fecundity selection, sometimes at the expense of viability. Ecological selection

14007-477: The average person to decode conversational implicature of emotional prosody has been found to be slightly less accurate than traditional facial expression discrimination ability; however, specific ability to decode varies by emotion. These emotional have been determined to be ubiquitous across cultures, as they are utilized and understood across cultures. Various emotions, and their general experimental identification rates, are as follows: The prosody of an utterance

14168-450: The best chance of being preserved in the struggle for life; and from the strong principle of inheritance they will tend to produce offspring similarly characterised. This principle of preservation, I have called, for the sake of brevity, Natural Selection. Once he had his theory, Darwin was meticulous about gathering and refining evidence before making his idea public. He was in the process of writing his "big book" to present his research when

14329-463: The children, Ninio also found that the pattern of subjects, direct objects and indirect objects in mothers’ speech does not provide the required prototypical semantics for the construction to be established. Adele Goldberg and her associates had previously reported similar negative results concerning the pattern of direct objects in parental speech. These findings are a blow to the CxG theory that relies on

14490-605: The cognitive semantics of communication—the modern version of the old Ross -Lakoff performative hypothesis from the 1960s. The form and content are symbolically linked in the sense advocated by Langacker . Thus a construction is treated like a sign in which all structural aspects are integrated parts and not distributed over different modules as they are in the componential model. Consequentially, not only constructions that are lexically fixed, like many idioms, but also more abstract ones like argument structure schemata, are pairings of form and conventionalized meaning. For instance,

14651-547: The componential model, construction grammar denies any strict distinction between the two and proposes a syntax–lexicon continuum . The argument goes that words and complex constructions are both pairs of form and meaning and differ only in internal symbolic complexity. Instead of being discrete modules and thus subject to very different processes they form the extremes of a continuum (from regular to idiosyncratic): syntax > subcategorization frame > idiom > morphology > syntactic category > word/ lexicon (these are

14812-476: The concept of the "cost" of natural selection. Sewall Wright elucidated the nature of selection and adaptation. In his book Genetics and the Origin of Species (1937), Theodosius Dobzhansky established the idea that mutation, once seen as a rival to selection, actually supplied the raw material for natural selection by creating genetic diversity. Ernst Mayr recognised the key importance of reproductive isolation for speciation in his Systematics and

14973-434: The different alleles, or variants of the gene that produces the variants of the trait. Selection can be divided into three classes, on the basis of its effect on allele frequencies: directional , stabilizing , and disruptive selection . Directional selection occurs when an allele has a greater fitness than others, so that it increases in frequency, gaining an increasing share in the population. This process can continue until

15134-458: The dog-days, but only if we have it in winter. If then, it is agreed that things are either the result of coincidence or for an end, and these cannot be the result of coincidence or spontaneity, it follows that they must be for an end; and that such things are all due to nature even the champions of the theory which is before us would agree. Therefore action for an end is present in things which come to be and are by nature. The struggle for existence

15295-579: The evolution of particular populations of a species, and populations that evolve to be sufficiently different eventually become different species. Darwin's ideas were inspired by the observations that he had made on the second voyage of HMS Beagle (1831–1836), and by the work of a political economist, Thomas Robert Malthus , who, in An Essay on the Principle of Population (1798), noted that population (if unchecked) increases exponentially , whereas

15456-399: The extravagant plumage of some male birds such as the peacock. An alternate theory proposed by the same Ronald Fisher in 1930 is the sexy son hypothesis , that mothers want promiscuous sons to give them large numbers of grandchildren and so choose promiscuous fathers for their children. Aggression between members of the same sex is sometimes associated with very distinctive features, such as

15617-512: The father of modern genetics, was a contemporary of Darwin's, his work lay in obscurity, only being rediscovered in 1900. With the early 20th-century integration of evolution with Mendel's laws of inheritance, the so-called modern synthesis , scientists generally came to accept natural selection. The synthesis grew from advances in different fields. Ronald Fisher developed the required mathematical language and wrote The Genetical Theory of Natural Selection (1930). J. B. S. Haldane introduced

15778-473: The fittest" does not give an "improvement in fitness", it only represents the removal of the less fit variants from a population. A mathematical example of "survival of the fittest" is given by Haldane in his paper "The Cost of Natural Selection". Haldane called this process "substitution" or more commonly in biology, this is called "fixation". This is correctly described by the differential survival and reproduction of individuals due to differences in phenotype. On

15939-412: The floor, to yield the turn, to invite a backchannel like uh-huh, and so on), and marking topic structure (starting a new topic, closing a topic, interpolating a parenthetical remark, and so on), among others. For example, David Brazil and his associates studied how intonation can indicate whether information is new or already established; whether a speaker is dominant or not in a conversation; and when

16100-472: The food supply grows only arithmetically ; thus, inevitable limitations of resources would have demographic implications, leading to a "struggle for existence". When Darwin read Malthus in 1838 he was already primed by his work as a naturalist to appreciate the "struggle for existence" in nature. It struck him that as population outgrew resources, "favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be

16261-420: The form of utterance (statement, question, or command), the presence of irony or sarcasm , certain emphasis on words or morphemes, contrast , focus , and so on. Prosody displays elements of language that are not encoded by grammar , punctuation or choice of vocabulary . In the study of prosodic aspects of speech, it is usual to distinguish between auditory measures ( subjective impressions produced in

16422-589: The formal aspects of constructions and makes use of a unification-based framework for description of syntax, not unlike head-driven phrase structure grammar . Its proponents/developers include Charles Fillmore, Paul Kay, Laura Michaelis , and to a certain extent Ivan Sag . Immanent within BCG works like Fillmore and Kay 1995 and Michaelis and Ruppenhofer 2001 is the notion that phrasal representations—embedding relations—should not be used to represent combinatoric properties of lexemes or lexeme classes. For example, BCG abandons

16583-421: The formation of new species." Darwin wrote: If during the long course of ages and under varying conditions of life, organic beings vary at all in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the high geometrical powers of increase of each species, at some age, season, or year, a severe struggle for life, and this certainly cannot be disputed; then, considering

16744-453: The function of many other genes. Most, but not all, mutations in regulatory genes result in non-viable embryos. Some nonlethal regulatory mutations occur in HOX genes in humans, which can result in a cervical rib or polydactyly , an increase in the number of fingers or toes. When such mutations result in a higher fitness, natural selection favours these phenotypes and the novel trait spreads in

16905-408: The highlighting of particular words to create different intonation patterns can be seen in the following English conversation: The exchange above is an example of using intonation to highlight particular words and to employ rising and falling of pitch to change meaning. If read out loud, the pitch of the voice moves in different directions on the word "cat." In the first line, pitch goes up, indicating

17066-641: The idea that form is semantically motivated. Embodied construction grammar (ECG), which is being developed by the Neural Theory of Language (NTL) group at ICSI, UC Berkeley, and the University of Hawaiʻi, particularly including Benjamin Bergen and Nancy Chang, adopts the basic constructionist definition of a grammatical construction, but emphasizes the relation of constructional semantic content to embodiment and sensorimotor experiences. A central claim

17227-487: The idea that tiny, virtually imperceptible changes in successive generations could produce consequences on the scale of differences between species. The early 19th-century zoologist Jean-Baptiste Lamarck suggested the inheritance of acquired characteristics as a mechanism for evolutionary change; adaptive traits acquired by an organism during its lifetime could be inherited by that organism's progeny, eventually causing transmutation of species . This theory, Lamarckism ,

17388-402: The identification and discrimination of semantically neutral sentences with varying tones of happiness, sadness, anger, and indifference, exemplifying the importance of prosody in language comprehension and production. Producing these nonverbal elements requires intact motor areas of the face, mouth, tongue, and throat. This area is associated with Brodmann areas 44 and 45 ( Broca's area ) of

17549-464: The indirect-object in VI. Anat Ninio examined the speech of a large sample of young English-speaking children and found that they do not in fact learn the syntactic patterns with the prototypical semantics claimed to be associated with them, or with any single semantics. The major reason is that such pairings are not consistently modelled for them in parental speech. Examining the maternal speech addressed to

17710-502: The infinite complexity of the relations of all organic beings to each other and to their conditions of existence, causing an infinite diversity in structure, constitution, and habits, to be advantageous to them, I think it would be a most extraordinary fact if no variation ever had occurred useful to each being's own welfare, in the same way as so many variations have occurred useful to man. But if variations useful to any organic being do occur, assuredly individuals thus characterised will have

17871-688: The internal structure of constructions. Radical construction grammar is totally non- reductionist , and Croft argues that constructions are not derived from their parts, but that the parts are derived from the constructions they appear in. Thus, in radical construction grammar, constructions are linked to Gestalts . Radical construction grammar rejects the idea that syntactic categories, roles, and relations are universal and argues that they are not only language-specific, but also construction specific. Thus, there are no universals that make reference to formal categories, since formal categories are language- and construction-specific. The only universals are to be found in

18032-416: The left frontal lobe . Damage to areas 44/45, specifically on the right hemisphere, produces motor aprosodia, with the nonverbal elements of speech being disturbed (facial expression, tone, rhythm of voice). Understanding these nonverbal elements requires an intact and properly functioning right-hemisphere perisylvian area , particularly Brodmann area 22 (not to be confused with the corresponding area in

18193-444: The level or unit of selection . Individual selection acts on the individual, in the sense that adaptations are "for" the benefit of the individual, and result from selection among individuals. Gene selection acts directly at the level of the gene. In kin selection and intragenomic conflict , gene-level selection provides a more apt explanation of the underlying process. Group selection , if it occurs, acts on groups of organisms, on

18354-526: The linguistic functions of intonation and stress, as well as other prosodic features such as rhythm and tempo. Additional prosodic variables have been studied, including voice quality and pausing. The behavior of the prosodic variables can be studied either as contours across the prosodic unit or by the behavior of boundaries. Prosodic features are suprasegmental, since they are properties of units of speech that are defined over groups of sounds rather than single segments. When talking about prosodic features, it

18515-403: The maintenance of the status quo by eliminating less fit variants. Selection can be classified in several different ways, such as by its effect on a trait, on genetic diversity, by the life cycle stage where it acts, by the unit of selection, or by the resource being competed for. Selection has different effects on traits. Stabilizing selection acts to hold a trait at a stable optimum, and in

18676-521: The meaning of an expression was not simply a function of the meanings of its parts. Instead, he suggested, constructions themselves must have meanings. Another early study was "There-Constructions," which appeared as Case Study 3 in George Lakoff's Women, Fire, and Dangerous Things . It argued that the meaning of the whole was not a function of the meanings of the parts, that odd grammatical properties of Deictic There-constructions followed from

18837-557: The mind of the listener) and objective measures (physical properties of the sound wave and physiological characteristics of articulation that may be measured objectively). Auditory (subjective) and objective ( acoustic and articulatory) measures of prosody do not correspond in a linear way. Most studies of prosody have been based on auditory analysis using auditory scales. Auditorily, the major prosodic variables are: Acoustically, these prosodic variables correspond closely to: Different combinations of these variables are exploited in

18998-406: The mutations that caused the maladaptation. At the same time, new mutations occur, resulting in a mutation–selection balance . The exact outcome of the two processes depends both on the rate at which new mutations occur and on the strength of the natural selection, which is a function of how unfavourable the mutation proves to be. Genetic linkage occurs when the loci of two alleles are close on

19159-546: The naturalist Alfred Russel Wallace independently conceived of the principle and described it in an essay he sent to Darwin to forward to Charles Lyell . Lyell and Joseph Dalton Hooker decided to present his essay together with unpublished writings that Darwin had sent to fellow naturalists, and On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection

19320-410: The opposite of over-dominance), and can result in a bimodal distribution of trait values. Finally, balancing selection can occur through frequency-dependent selection, where the fitness of one particular phenotype depends on the distribution of other phenotypes in the population. The principles of game theory have been applied to understand the fitness distributions in these situations, particularly in

19481-412: The other hand, "improvement in fitness" can occur in an environment where "survival of the fittest" is also acting. Richard Lenski 's classic E. coli long-term evolution experiment is an example of adaptation in a competitive environment, ("improvement in fitness" during "survival of the fittest"). The probability of a beneficial mutation occurring on some member of the lineage to give improved fitness

19642-405: The other hand, "improvement in fitness" is not dependent on the differential survival and reproduction of individuals due to differences in phenotype, it is dependent on the absolute survival of the particular variant. The probability of a beneficial mutation occurring on some member of a population depends on the total number of replications of that variant. The mathematics of "improvement in fitness

19803-406: The patterns concerning the mapping of meaning onto form. Radical construction grammar rejects the notion of syntactic relations altogether and replaces them with semantic relations. Like Goldbergian/Lakovian construction grammar and cognitive grammar, radical construction grammar is closely related to cognitive linguistics, and like cognitive grammar, radical construction grammar appears to be based on

19964-449: The perception of word groups, or chunks. Examples include the phrase , phraseme , constituent or interjection . Chunks commonly highlight lexical items or fixed expression idioms . Chunking prosody is present on any complete utterance and may correspond to a syntactic category , but not necessarily. The well-known English chunk "Know what I mean?" in common usage sounds like a single word ("No-wada-MEEN?") due to blurring or rushing

20125-404: The phenotype. Some traits are governed by only a single gene, but most traits are influenced by the interactions of many genes. A variation in one of the many genes that contributes to a trait may have only a small effect on the phenotype; together, these genes can produce a continuum of possible phenotypic values. When some component of a trait is heritable, selection alters the frequencies of

20286-511: The phrase survival of the fittest , which became a popular summary of the theory. The fifth edition of On the Origin of Species published in 1869 included Spencer's phrase as an alternative to natural selection, with credit given: "But the expression often used by Mr. Herbert Spencer of the Survival of the Fittest is more accurate, and is sometimes equally convenient." Although the phrase

20447-441: The phrase "either invariably or normally come about", not the origin of species: ... Yet it is impossible that this should be the true view. For teeth and all other natural things either invariably or normally come about in a given way; but of not one of the results of chance or spontaneity is this true. We do not ascribe to chance or mere coincidence the frequency of rain in winter, but frequent rain in summer we do; nor heat in

20608-688: The point that the methicillin-resistant Staphylococcus aureus (MRSA) has been described as a "superbug" because of the threat it poses to health and its relative invulnerability to existing drugs. Response strategies typically include the use of different, stronger antibiotics; however, new strains of MRSA have recently emerged that are resistant even to these drugs. This is an evolutionary arms race , in which bacteria develop strains less susceptible to antibiotics, while medical researchers attempt to develop new antibiotics that can kill them. A similar situation occurs with pesticide resistance in plants and insects. Arms races are not necessarily induced by man;

20769-406: The population. Established traits are not immutable; traits that have high fitness in one environmental context may be much less fit if environmental conditions change. In the absence of natural selection to preserve such a trait, it becomes more variable and deteriorate over time, possibly resulting in a vestigial manifestation of the trait, also called evolutionary baggage . In many circumstances,

20930-563: The population. In this way the natural environment of an organism "selects for" traits that confer a reproductive advantage, causing evolutionary change, as Darwin described. This gives the appearance of purpose, but in natural selection there is no intentional choice. Artificial selection is purposive where natural selection is not, though biologists often use teleological language to describe it. The peppered moth exists in both light and dark colours in Great Britain, but during

21091-421: The pragmatic meaning of the construction, and that variations on the central construction could be seen as simple extensions using form-meaning pairs of the central construction. Fillmore et al.'s (1988) paper on the English let alone construction was a second classic. These two papers propelled cognitive linguists into the study of CxG. Since the late 1990s there has been a shift towards a general preference for

21252-551: The proverbial expression The bigger they come, the harder they fall . Construction grammarians point out that this is not merely a fixed phrase; the Correlative Conditional is a general pattern ( The Xer, the Yer ) with "slots" that can be filled by almost any comparative phrase (e.g. The more you think about it, the less you understand ). Advocates of CxG argue these kinds of idiosyncratic patterns are more common than

21413-527: The rate of reproduction, given survival. Others split the life cycle into further components of selection. Thus viability and survival selection may be defined separately and respectively as acting to improve the probability of survival before and after reproductive age is reached, while fecundity selection may be split into additional sub-components including sexual selection, gametic selection, acting on gamete survival, and compatibility selection, acting on zygote formation. Selection can also be classified by

21574-469: The recognition of emotion may be quite low, of the order of 50%, hampering the complex interrelationship function of speech advocated by some authors. However, even if emotional expression through prosody cannot always be consciously recognized, tone of voice may continue to have subconscious effects in conversation. This sort of expression stems not from linguistic or semantic effects, and can thus be isolated from traditional linguistic content. Aptitude of

21735-403: The same as memetics. Lastly, the most basic syntactic patterns of English, namely the core grammatical relations subject-verb, verb object and verb-indirect object, are counter-evidence for the very concept of constructions as pairings of linguistic patterns with meanings. Instead of the postulated form-meaning pairing, core grammatical relations possess a wide variability of semantics, exhibiting

21896-411: The same proposition are not synonymous, but display differences in content: in this case the pragmatic content. As mentioned above, Construction grammar is a "family" of theories rather than one unified theory. There are a number of formalized Construction grammar frameworks. Some of these are: Berkeley Construction Grammar (BCG: formerly also called simply Construction Grammar in upper case) focuses on

22057-421: The same sex, that is male–male competition, or intersexual , where one gender chooses mates , most often with males displaying and females choosing. However, in some species, mate choice is primarily by males, as in some fishes of the family Syngnathidae . Phenotypic traits can be displayed in one sex and desired in the other sex, causing a positive feedback loop called a Fisherian runaway , for example,

22218-427: The second syllable. For example: Here, adults will emphasize the first syllable, "IN", as "increase" functions as an adjective. Here, adults will emphasize the second syllable, "CREASE", as "increase" functions as a verb. Another way that lexical prosody is used in the English language is in compound nouns such as "wishbone, mailbox, and blackbird" where the first compound is emphasized. Some suffixes can also affect

22379-469: The selection of individuals that reproduce", without regard to whether the basis of the selection is heritable. Traits that cause greater reproductive success of an organism are said to be selected for , while those that reduce success are selected against . Natural variation occurs among the individuals of any population of organisms. Some differences may improve an individual's chances of surviving and reproducing such that its lifetime reproductive rate

22540-496: The sentence into chunks ) and changes in intonation will reduce or remove the ambiguity. Moving the intonational boundary in cases such as the above example will tend to change the interpretation of the sentence. This result has been found in studies performed in both English and Bulgarian. Research in English word recognition has demonstrated an important role for prosody. Intonation and stress work together to highlight important words or syllables for contrast and focus . This

22701-444: The simplest case all deviations from this optimum are selectively disadvantageous. Directional selection favours extreme values of a trait. The uncommon disruptive selection also acts during transition periods when the current mode is sub-optimal, but alters the trait in more than one direction. In particular, if the trait is quantitative and univariate then both higher and lower trait levels are favoured. Disruptive selection can be

22862-426: The speaker's pitch level returns to the level typical of the onset of a new intonation unit. In this way potential ambiguities may be resolved. For example, the sentence "They invited Bob and Bill and Al got rejected" is ambiguous when written, although addition of a written comma after either "Bob" or "Bill" will remove the sentence's ambiguity. But when the sentence is read aloud, prosodic cues like pauses (dividing

23023-494: The study of kin selection and the evolution of reciprocal altruism . A portion of all genetic variation is functionally neutral, producing no phenotypic effect or significant difference in fitness. Motoo Kimura 's neutral theory of molecular evolution by genetic drift proposes that this variation accounts for a large fraction of observed genetic diversity. Neutral events can radically reduce genetic variation through population bottlenecks . which among other things can cause

23184-407: The term "Natural Selection", preferring the term "Natural Preservation". For Darwin and his contemporaries, natural selection was in essence synonymous with evolution by natural selection. After the publication of On the Origin of Species , educated people generally accepted that evolution had occurred in some form. However, natural selection remained controversial as a mechanism, partly because it

23345-430: The term "natural selection", contrasting it with artificial selection , which is intentional, whereas natural selection is not. Variation of traits, both genotypic and phenotypic , exists within all populations of organisms . However, some traits are more likely to facilitate survival and reproductive success . Thus, these traits are passed onto the next generation. These traits can also become more common within

23506-482: The theory itself and a variety of applications can be found in Jerome Feldman's From Molecule to Metaphor: A Neural Theory of Language (MIT Press, 2006). Fluid construction grammar (FCG) was designed by Luc Steels and his collaborators for doing experiments on the origins and evolution of language . FCG is a fully operational and computationally implemented formalism for construction grammars and proposes

23667-477: The time, anger 95%, surprise 91%, sadness 81%, and neutral tone 76%. When a database of this speech was processed by computer, segmental features allowed better than 90% recognition of happiness and anger, while suprasegmental prosodic features allowed only 44%–49% recognition. The reverse was true for surprise, which was recognized only 69% of the time by segmental features and 96% of the time by suprasegmental prosody. In typical conversation (no actor voice involved),

23828-441: The timing of successive units of speech, a regularity referred to as isochrony , and that every language may be assigned one of three rhythmical types: stress-timed (where the durations of the intervals between stressed syllables is relatively constant), syllable-timed (where the durations of successive syllables are relatively constant) and mora-timed (where the durations of successive morae are relatively constant). As explained in

23989-644: The traditional practice of using non-branching domination (NP over N' over N) to describe undetermined nominals that function as NPs, instead introducing a determination construction that requires ('asks for') a non-maximal nominal sister and a lexical 'maximality' feature for which plural and mass nouns are unmarked. BCG also offers a unification-based representation of 'argument structure' patterns as abstract verbal lexeme entries ('linking constructions'). These linking constructions include transitive, oblique goal and passive constructions. These constructions describe classes of verbs that combine with phrasal constructions like

24150-447: The traditional terms; construction grammars use a different terminology). In construction grammar, the grammar of a language is made up of taxonomic networks of families of constructions, which are based on the same principles as those of the conceptual categories known from cognitive linguistics , such as inheritance, prototypicality, extensions, and multiple parenting. Four different models are proposed in relation to how information

24311-445: The usage-based model. The shift towards the usage-based approach in construction grammar has inspired the development of several corpus -based methodologies of constructional analysis (for example, collostructional analysis ). One of the most distinctive features of CxG is its use of multi-word expressions and phrasal patterns as the building blocks of syntactic analysis. One example is the Correlative Conditional construction, found in

24472-518: The useful forms survived: So what hinders the different parts [of the body] from having this merely accidental relation in nature? as the teeth, for example, grow by necessity, the front ones sharp, adapted for dividing, and the grinders flat, and serviceable for masticating the food; since they were not made for the sake of this, but it was the result of accident. And in like manner as to the other parts in which there appears to exist an adaptation to an end. Wheresoever, therefore, all things together (that

24633-447: The ways in which different words are stressed. Take "active" for example. Without the suffix, the lexical emphasis is on "AC". However, when we add the suffix -ity, the stress shifts to "TIV". Phrasal prosody refers to the rhythm and tempo of phrases, often in an artistic setting such as music or poetry, but not always. The rhythm of the English language has four different elements: stress, time, pause, and pitch. Furthermore, "When stress

24794-420: The works of William Dwight Whitney , Leonard Bloomfield , Charles Hockett , and others. At the same time, the claim made by construction grammarians, that their research represents a continuation of Saussurean linguistics, has been considered misleading. German philologist Elisabeth Leiss regards construction grammar as regress, linking it with the 19th century social darwinism of August Schleicher . There

24955-421: Was achieved for a purpose, citing the regularity of heredity in species as proof. Nevertheless, he accepted in his biology that new types of animals, monstrosities (τερας), can occur in very rare instances ( Generation of Animals , Book IV). As quoted in Darwin's 1872 edition of The Origin of Species , Aristotle considered whether different forms (e.g., of teeth) might have appeared accidentally, but only

25116-611: Was an influence on the Soviet biologist Trofim Lysenko 's ill-fated antagonism to mainstream genetic theory as late as the mid-20th century. Between 1835 and 1837, the zoologist Edward Blyth worked on the area of variation, artificial selection, and how a similar process occurs in nature. Darwin acknowledged Blyth's ideas in the first chapter on variation of On the Origin of Species . In 1859, Charles Darwin set out his theory of evolution by natural selection as an explanation for adaptation and speciation. He defined natural selection as

25277-482: Was described by Kleinman. An empirical example of "improvement in fitness" is given by the Kishony Mega-plate experiment. In this experiment, "improvement in fitness" depends on the number of replications of the particular variant for a new variant to appear that is capable of growing in the next higher drug concentration region. Fixation or substitution is not required for this "improvement in fitness". On

25438-493: Was later described by the Islamic writer Al-Jahiz in the 9th century, particularly in the context of top-down population regulation, but not in reference to individual variation or natural selection. At the turn of the 16th century Leonardo da Vinci collected a set of fossils of ammonites as well as other biological material. He extensively reasoned in his writings that the shapes of animals are not given once and forever by

25599-429: Was perceived to be too weak to explain the range of observed characteristics of living organisms, and partly because even supporters of evolution balked at its "unguided" and non- progressive nature, a response that has been characterised as the single most significant impediment to the idea's acceptance. However, some thinkers enthusiastically embraced natural selection; after reading Darwin, Herbert Spencer introduced

25760-797: Was read to the Linnean Society of London announcing co-discovery of the principle in July 1858. Darwin published a detailed account of his evidence and conclusions in On the Origin of Species in 1859. In the 3rd edition of 1861 Darwin acknowledged that others—like William Charles Wells in 1813, and Patrick Matthew in 1831—had proposed similar ideas, but had neither developed them nor presented them in notable scientific publications. Darwin thought of natural selection by analogy to how farmers select crops or livestock for breeding, which he called " artificial selection "; in his early manuscripts he referred to

25921-487: Was reversed by the effect of the Clean Air Act 1956 , and the dark moths became rare again, demonstrating the influence of natural selection on peppered moth evolution . A recent study, using image analysis and avian vision models, shows that pale individuals more closely match lichen backgrounds than dark morphs and for the first time quantifies the camouflage of moths to predation risk. The concept of fitness

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