A Medley of Potpourri: May 28, 2015

Thursday, May 28, 2015

Classical electromagnetism and special relativity

From Wikipedia, the free encyclopedia

The theory of special relativity plays an important role in the modern theory of classical electromagnetism. First of all, it gives formulas for how electromagnetic objects, in particular the electric and magnetic fields, are altered under a Lorentz transformation from one inertial frame of reference to another. Secondly, it sheds light on the relationship between electricity and magnetism, showing that frame of reference determines if an observation follows electrostatic or magnetic laws. Third, it motivates a compact and convenient notation for the laws of electromagnetism, namely the "manifestly covariant" tensor form.

Maxwell's equations, when they were first stated in their complete form in 1865, would turn out to be compatible with special relativity.^[1] Moreover, the apparent coincidences in which the same effect was observed due to different physical phenomena by two different observers would be shown to be not coincidental in the least by special relativity. In fact, half of Einstein's 1905 first paper on special relativity, "On the Electrodynamics of Moving Bodies," explains how to transform Maxwell's equations.

Transformation of the fields between inertial frames

The E and B fields

This equation, also called the Joules-Bernoulli equation, considers two inertial frames. As notation, the field variables in one frame are unprimed, and in a frame moving relative to the unprimed frame at velocity v, the fields are denoted with primes. In addition, the fields parallel to the velocity v are denoted by

\stackrel{\mathbf {E}_{\parallel}}{}

while the fields perpendicular to v are denoted as

\stackrel{\mathbf {E}_{\bot}}{}

. In these two frames moving at relative velocity v, the E-fields and B-fields are related by:^[2]

\begin{align} & \mathbf {{E}_{\parallel}}' = \mathbf {{E}_{\parallel}}\\ & \mathbf {{B}_{\parallel}}' = \mathbf {{B}_{\parallel}}\\ & \mathbf {{E}_{\bot}}'= \gamma \left( \mathbf {E}_{\bot} + \mathbf{ v} \times \mathbf {B} \right) \\ & \mathbf {{B}_{\bot}}'= \gamma \left( \mathbf {B}_{\bot} -\frac{1}{c^2} \mathbf{ v} \times \mathbf {E} \right) \end{align}

where

\gamma \ \overset{\underset{\mathrm{def}}{}}{=} \ \frac{1}{\sqrt{1 - v^2/{c}^2}}

is called the Lorentz factor and c is the speed of light in free space. The inverse transformations are the same except v → −v.

An equivalent, alternative expression is:^[3]

\begin{align} & \mathbf{E}' = \gamma \left( \mathbf{E} + \mathbf{v} \times \mathbf{B} \right ) - \left ({\gamma-1} \right ) ( \mathbf{E} \cdot \mathbf{\hat{v}} ) \mathbf{\hat{v}} \\ & \mathbf{B}' = \gamma \left( \mathbf{B} - \frac {\mathbf{v} \times \mathbf{E}}{c^2} \right ) - \left ({\gamma-1} \right ) ( \mathbf{B} \cdot \mathbf{\hat{v}} ) \mathbf{\hat{v}}\\ \end{align}

where v̂ is the velocity unit vector.

If one of the fields is zero in one frame of reference, that doesn't necessarily mean it is zero in all other frames of reference. This can be seen by, for instance, making the unprimed electric field zero in the transformation to the primed electric field. In this case, depending on the orientation of the magnetic field, the primed system could see an electric field, even though there is none in the unprimed system.

This does not mean two completely different sets of events are seen in the two frames, but that the same sequence of events is described in two different ways (see Moving magnet and conductor problem below).

If a particle of charge q moves with velocity u with respect to frame S, then the Lorentz force in frame S is:

\mathbf{F}=q\mathbf{E}+q \mathbf{u} \times \mathbf{B}

In frame S', the Lorentz force is:

\mathbf{F'}=q\mathbf{E'}+q \mathbf{u'} \times \mathbf{B'}

If S and S' have aligned axes then:^[4]

\begin{align} & u_x'=\frac{u_x+v}{1 + (v \ u_x)/c^2}\\ & u_y'=\frac{u_y/\gamma}{1 + (v \ u_x)/c^2}\\ & u_z'=\frac{u_z/\gamma}{1 + (v \ u_x)/c^2} \end{align}

A derivation for the transformation of the Lorentz force for the particular case u = 0 is given here.^[5] A more general one can be seen here.^[6]

Component by component, for relative motion along the x-axis, this works out to be the following:

\begin{align} & E'_x = E_x & \qquad & B'_x = B_x \\ & E'_y = \gamma \left( E_y - v B_z \right) & & B'_y = \gamma \left( B_y + \frac{v}{c^2} E_z \right) \\ & E'_z = \gamma \left( E_z + v B_y \right) & & B'_z = \gamma \left( B_z - \frac{v}{c^2} E_y \right). \\ \end{align}

The transformations in this form can be made more compact by introducing the electromagnetic tensor (defined below), which is a covariant tensor.

The D and H fields

For the electric displacement D and magnetic intensity H, using the constitutive relations and the result for c²:

\mathbf{D}=\epsilon_0\mathbf{E}\,, \quad \mathbf{B}=\mu_0\mathbf{H}\,,\quad c^2=\frac{1}{\epsilon_0\mu_0}\,,

gives

\begin{align} \mathbf{D}' & =\gamma \left( \mathbf{D}+\frac{1}{c^2}\mathbf{v}\times \mathbf{H} \right)+(1-\gamma )(\mathbf{D}\cdot \mathbf{\hat{v}})\mathbf{\hat{v}} \\ \mathbf{H}' & =\gamma \left( \mathbf{H}-\mathbf{v}\times \mathbf{D} \right)+(1-\gamma )(\mathbf{H}\cdot \mathbf{\hat{v}})\mathbf{\hat{v}} \\ \end{align}

Analogously for E and B, the D and H form the electromagnetic displacement tensor.

The φ and A fields

An alternative simpler transformation of the EM field uses the electromagnetic potentials - the electric potential φ and magnetic potential A:^[7]

\begin{align} & \varphi' = \gamma (\varphi - v A_\parallel)\\ & A_\parallel' = \gamma (A_\parallel - v \varphi /c^2)\\ & A_\bot' = A_\bot \end{align}

where

\scriptstyle A_\parallel

is the parallel component of A to the direction of relative velocity between frames v, and

\scriptstyle A_\bot

is the perpendicular component. These transparently resemble the characteristic form of other Lorentz transformations (like time-position and energy-momentum), while the transformations of E and B above are slightly more complicated. The components can be collected together as:

\begin{align} \mathbf{A}' & = \mathbf{A} - \dfrac{\gamma \varphi}{c^2}\mathbf{v} + (\gamma-1) (\mathbf{A}\cdot\mathbf{\hat{v}})\mathbf{\hat{v}} \\ {\varphi}' & =\gamma \left( \varphi - \mathbf{A}\cdot \mathbf{v} \right) \end{align}

The ρ and J fields

Analogously for the charge density ρ and current density J,^[7]

\begin{align} & J_\parallel' = \gamma ( J_\parallel - v\rho)\\ & \rho' = \gamma (\rho - v J_\parallel /c^2)\\ & J_\bot' = J_\bot \end{align}

Collecting components together:

\begin{align} \mathbf{J}' & =\mathbf{J}-\gamma \rho \mathbf{v} +\left( \gamma -1 \right)(\mathbf{J}\cdot \mathbf{\hat{v}})\mathbf{\hat{v}} \\ {\rho }' & =\gamma ( \rho - \mathbf{J}\cdot \mathbf{v}/c^2) \end{align}

Non-relativistic approximations

For speeds v ≪ c, the relativistic factor γ ≈ 1, which yields:

\begin{align} \mathbf{E}' & \approx \mathbf{E}+\mathbf{v}\times \mathbf{B} \\ \mathbf{B}' & \approx \mathbf{B}-\frac{1}{c^2}\mathbf{v}\times \mathbf{E} \\ \mathbf{J}' & \approx \mathbf{J}-\rho \mathbf{v}\\ \rho' & \approx \left( \rho -\frac{1}{c^2}\mathbf{j}\cdot \mathbf{v} \right) \end{align}

so that there is no need to distinguish between the spatial and temporal coordinates in Maxwell's equations.

Relationship between electricity and magnetism

“

One part of the force between moving charges we call the magnetic force. It is really one aspect of an electrical effect.

”

—Richard Feynman^[8]

Deriving magnetism from electrostatics

The chosen reference frame determines if an electromagnetic phenomenon is viewed as an effect of electrostatics or magnetism. Authors usually derive magnetism from electrostatics when special relativity and charge invariance are taken into account. The Feynman Lectures on Physics (vol. 2, ch. 13-6) uses this method to derive the "magnetic" force on a moving charge next to a current-carrying wire. See also Haskell^[9] and Landau.^[10]

Fields intermix in different frames

The above transformation rules show that the electric field in one frame contributes to the magnetic field in another frame, and vice versa.^[11] This is often described by saying that the electric field and magnetic field are two interrelated aspects of a single object, called the electromagnetic field. Indeed, the entire electromagnetic field can be encoded in a single rank-2 tensor called the electromagnetic tensor; see below.

Moving magnet and conductor problem

A famous example of the intermixing of electric and magnetic phenomena in different frames of reference is called the "moving magnet and conductor problem", cited by Einstein in his 1905 paper on Special Relativity.
If a conductor moves with a constant velocity through the field of a stationary magnet, eddy currents will be produced due to a magnetic force on the electrons in the conductor. In the rest frame of the conductor, on the other hand, the magnet will be moving and the conductor stationary. Classical electromagnetic theory predicts that precisely the same microscopic eddy currents will be produced, but they will be due to an electric force.^[12]

Covariant formulation in vacuum

The laws and mathematical objects in classical electromagnetism can be written in a form which is manifestly covariant. Here, this is only done so for vacuum (or for the microscopic Maxwell equations, not using macroscopic descriptions of materials such as electric permittivity), and uses SI units.

This section uses Einstein notation, including Einstein summation convention. See also Ricci calculus for a summary of tensor index notations, and raising and lowering indices for definition of superscript and subscript indices, and how to switch between them. The Minkowski metric tensor η here has metric signature (+ − − −).

Field tensor and 4-current

The above relativistic transformations suggest the electric and magnetic fields are coupled together, in a mathematical object with 6 components: an antisymmetric second-rank tensor, or a bivector. This is called the electromagnetic field tensor, usually written as F^μν. In matrix form:^[13]

F^{\mu \nu} = \begin{pmatrix} 0 & -E_x/c & -E_y/c & -E_z/c \\ E_x/c & 0 & -B_z & B_y \\ E_y/c & B_z & 0 & -B_x \\ E_z/c & -B_y & B_x & 0 \end{pmatrix}

where c the speed of light - in natural units c = 1.

There is another way of merging the electric and magnetic fields into an antisymmetric tensor, by replacing E/c → B and B → − E/c, to get the dual tensor G^μν.

G^{\mu \nu} = \begin{pmatrix} 0 & -B_x & -B_y & -B_z \\ B_x & 0 & E_z/c & - E_y/c \\ B_y & -E_z/c & 0 & E_x/c \\ B_z & E_y/c & -E_x/c & 0 \end{pmatrix}

In the context of special relativity, both of these transform according to the Lorentz transformation according to

F'^{\alpha \beta} = \Lambda^\alpha{}_\mu \Lambda^\beta{}_\nu F^{\mu \nu}

where Λ^α_ν is the Lorentz transformation tensor for a change from one reference frame to another. The same tensor is used twice in the summation.

The charge and current density, the sources of the fields, also combine into the four-vector

J^\alpha = \begin{pmatrix} c \rho & J_x & J_y & J_z \end{pmatrix}

called the four-current.

Maxwell's equations in tensor form

Using these tensors, Maxwell's equations reduce to:^[13]

Maxwell's equations (Covariant formulation)

\begin{align} & \frac{\partial F^{\alpha \beta}}{\partial x^\alpha} = \mu_0 J^\beta\\ & \frac{\partial G^{\alpha \beta}}{\partial x^\alpha} = 0 \end{align}

where the partial derivatives may be written in various ways, see 4-gradient. The first equation listed above corresponds to both Gauss's Law (for β = 0) and the Ampère-Maxwell Law (for β = 1, 2, 3). The second equation corresponds to the two remaining equations, Gauss's law for magnetism (for β = 0) and Faraday's Law ( for β = 1, 2, 3).

These tensor equations are manifestly-covariant, meaning the equations can be seen to be covariant by the index positions. This short form of writing Maxwell's equations illustrates an idea shared amongst some physicists, namely that the laws of physics take on a simpler form when written using tensors.

By lowering the indices on F^αβ to obtain F_αβ (see raising and lowering indices):

F_{\alpha\beta} = \eta_{\alpha\lambda} \eta_{\beta\mu} F^{\lambda\mu}

the second equation can be written in terms of F_αβ as:

\epsilon^{\delta\alpha\beta\gamma} \dfrac{\partial F_{\beta\gamma}}{\partial x^\alpha} = \dfrac{\partial F_{\alpha\beta}}{\partial x^\gamma} + \dfrac{\partial F_{\gamma\alpha}}{\partial x^\beta} + \dfrac{\partial F_{\beta\gamma}}{\partial x^\alpha} = 0

where

\epsilon^{\alpha\beta\gamma\delta}

is the contravariant Levi-Civita symbol. Notice the cyclic permutation of indices in this equation:

\begin{array}{rc} & \scriptstyle{\alpha\,\, \longrightarrow \,\, \beta} \\ & \nwarrow_\gamma \swarrow \end{array}

.
Another covariant electromagnetic object is the electromagnetic stress-energy tensor, a covariant rank-2 tensor which includes the Poynting vector, Maxwell stress tensor, and electromagnetic energy density.

4-potential[edit]

The EM field tensor can also be written^[14]

F^{\alpha \beta} = \frac {\partial A^{\beta}}{\partial x_{\alpha}} - \frac {\partial A^{\alpha}}{\partial x_{\beta}} \, ,

where

A^\alpha = (\varphi/c, A_x,A_y,A_z)\,,

is the four-potential and

x_\alpha = (ct,-x,-y,-z )

is the four-position.

Using the 4-potential in the Lorenz gauge, an alternative manifestly-covariant formulation can be found in a single equation (a generalization of an equation due to Bernhard Riemann by Arnold Sommerfeld, known as the Riemann–Sommerfeld equation,^[15] or the covariant form of the Maxwell equations^[16]):

Maxwell's equations (Covariant Lorenz gauge formulation)

\Box A^\mu = \mu_0 J^\mu

where

\Box

is the d'Alembertian operator, or four-Laplacian. For a more comprehensive presentation of these topics, see Covariant formulation of classical electromagnetism.

How spacetime is built by quantum entanglement

A collaboration of physicists and a mathematician has made a significant step toward unifying general relativity and quantum mechanics by explaining how spacetime emerges from quantum entanglement in a more fundamental theory. The paper announcing the discovery by Hirosi Ooguri, a Principal Investigator at the University of Tokyo's Kavli IPMU, with Caltech mathematician Matilde Marcolli and graduate students Jennifer Lin and Bogdan Stoica, will be published in Physical Review Letters as an Editors' Suggestion "for the potential interest in the results presented and on the success of the paper in communicating its message, in particular to readers from other fields."

Physicists and mathematicians have long sought a Theory of Everything (ToE) that unifies general relativity and quantum mechanics. General relativity explains gravity and large-scale phenomena such as the dynamics of stars and galaxies in the universe, while quantum mechanics explains microscopic phenomena from the subatomic to molecular scales.

The holographic principle is widely regarded as an essential feature of a successful Theory of Everything. The holographic principle states that gravity in a three-dimensional volume can be described by quantum mechanics on a two-dimensional surface surrounding the volume. In particular, the three dimensions of the volume should emerge from the two dimensions of the surface. However, understanding the precise mechanics for the emergence of the volume from the surface has been elusive.

Now, Ooguri and his collaborators have found that quantum entanglement is the key to solving this question. Using a quantum theory (that does not include gravity), they showed how to compute energy density, which is a source of gravitational interactions in three dimensions, using quantum entanglement data on the surface. This is analogous to diagnosing conditions inside of your body by looking at X-ray images on two-dimensional sheets. This allowed them to interpret universal properties of quantum entanglement as conditions on the energy density that should be satisfied by any consistent quantum theory of gravity, without actually explicitly including gravity in the theory. The importance of quantum entanglement has been suggested before, but its precise role in emergence of spacetime was not clear until the new paper by Ooguri and collaborators.

Quantum entanglement is a phenomenon whereby quantum states such as spin or polarization of particles at different locations cannot be described independently. Measuring (and hence acting on) one particle must also act on the other, something that Einstein called "spooky action at distance." The work of Ooguri and collaborators shows that this quantum entanglement generates the extra dimensions of the gravitational theory.

"It was known that quantum entanglement is related to deep issues in the unification of general relativity and quantum mechanics, such as the black hole information paradox and the firewall paradox," says Hirosi Ooguri. "Our paper sheds new light on the relation between quantum entanglement and the microscopic structure of spacetime by explicit calculations. The interface between quantum gravity and information science is becoming increasingly important for both fields. I myself am collaborating with information scientists to pursue this line of research further."

More information: Locality of Gravitational Systems from Entanglement of Conformal Field Theories, Physical Review Letters, 2015.

Provided by University of Tokyo

Molecular evolution

From Wikipedia, the free encyclopedia

Molecular evolution is a change in the sequence composition of cellular molecules such as DNA, RNA, and proteins across generations. The field of molecular evolution uses principles of evolutionary biology and population genetics to explain patterns in these changes. Major topics in molecular evolution concern the rates and impacts of single nucleotide changes, neutral evolution vs. natural selection, origins of new genes, the genetic nature of complex traits, the genetic basis of speciation, evolution of development, and ways that evolutionary forces influence genomic and phenotypic changes.

Forces in molecular evolution

The content and structure of a genome is the product of the molecular and population genetic forces which act upon that genome. Novel genetic variants will arise through mutation and will spread and be maintained in populations due to genetic drift or natural selection.

Mutation

Mutations are permanent, transmissible changes to the genetic material (DNA or RNA) of a cell or virus. Mutations result from errors in DNA replication during cell division and by exposure to radiation, chemicals, and other environmental stressors, or viruses and transposable elements. Most mutations that occur are Single nucleotide polymorphisms which modify single bases of the DNA sequence. Other types of mutations modify larger segments of DNA and can cause duplications, insertions, deletions, inversions, and translocations.
Most organisms display a strong bias in the types of mutations that occur with strong influence in GC content. Transitions (A ↔ G or C ↔ T) are more common than transversions (purine ↔ pyrimidine)^[1] and are less likely to alter amino acid sequences of proteins.

Mutations are stochastic and typically occur randomly across genes. Mutation rates for single nucleotide sites for most organisms are very low, roughly 10⁻⁹ to 10⁻⁸ per site per generation, though some viruses have higher mutation rates on the order of 10⁻⁶ per site per generation. Among these mutations, some will be neutral or beneficial and will remain in the genome unless lost via Genetic drift, and others will be detrimental and will be eliminated from the genome by natural selection.

Because mutations are extremely rare, they accumulate very slowly across generations. While the number of mutations which appears in any single generation may vary, over very long time periods they will appear to accumulate at a regular pace. Using the mutation rate per generation and the number of nucleotide differences between two sequences, divergence times can be estimated effectively via the molecular clock.

Recombination

Recombination is a process that results in genetic exchange between chromosomes or chromosomal regions.
Recombination counteracts physical linkage between adjacent genes, thereby reducing genetic hitchhiking. The resulting independent inheritance of genes results in more efficient selection, meaning that regions with higher recombination will harbor fewer detrimental mutations, more selectively favored variants, and fewer errors in replication and repair. Recombination can also generate particular types of mutations if chromosomes are misaligned.

Gene conversion

Gene conversion is a type of recombination that is the product of DNA repair where nucleotide damage is corrected using orthologous genomic regions as a template. Damaged bases are first excised, the damaged strand is then aligned with an undamaged homolog, and DNA synthesis repairs the excised region using the undamaged strand as a guide. Gene conversion is often responsible for homogenizing sequence of duplicate genes over long time periods, reducing nucleotide divergence.

Genetic drift

Genetic drift is the change of allele frequencies from one generation to the next due to stochastic effects of random sampling in finite populations. Some existing variants have no effect on fitness and may increase or decrease in frequency simply due to chance. "Nearly neutral" variants whose selection coefficient is close to a threshold value of 1 / the effective population size will also be affected by chance as well as by selection and mutation. Many genomic features have been ascribed to accumulation of nearly neutral detrimental mutations as a result of small effective population sizes.^[2] With a smaller effective population size, a larger variety of mutations will behave as if they are neutral due to inefficiency of selection.

Selection

Selection occurs when organisms with greater fitness, i.e. greater ability to survive or reproduce, are favored in subsequent generations, thereby increasing the instance of underlying genetic variants in a population. Selection can be the product of natural selection, artificial selection, or sexual selection. Natural selection is any selective process that occurs due to the fitness of an organism to its environment. In contrast sexual selection is a product of mate choice and can favor the spread of genetic variants which act counter to natural selection but increase desirability to the opposite sex or increase mating success. Artificial selection, also known as selective breeding, is imposed by an outside entity, typically humans, in order to increase the frequency of desired traits.

The principles of population genetics apply similarly to all types of selection, though in fact each may produce distinct effects due to clustering of genes with different functions in different parts of the genome, or due to different properties of genes in particular functional classes. For instance, sexual selection could be more likely to affect molecular evolution of the sex chromosomes due to clustering of sex specific genes on the X,Y,Z or W.

Selection can operate at the gene level at the expense of organismal fitness, resulting in a selective advantage for selfish genetic elements in spite of a host cost. Examples of such selfish elements include transposable elements, meiotic drivers, killer X chromosomes, selfish mitochondria, and self-propagating introns. (See Intragenomic conflict.)

Genome architecture

Genome size

Genome size is influenced by the amount of repetitive DNA as well as number of genes in an organism. The C-value paradox refers to the lack of correlation between organism 'complexity' and genome size. Explanations for the so-called paradox are two-fold. First, repetitive genetic elements can comprise large portions of the genome for many organisms, thereby inflating DNA content of the haploid genome. Secondly, the number of genes is not necessarily indicative of the number of developmental stages or tissue types in an organism. An organism with few developmental stages or tissue types may have large numbers of genes that influence non-developmental phenotypes, inflating gene content relative to developmental gene families.

Neutral explanations for genome size suggest that when population sizes are small, many mutations become nearly neutral. Hence, in small populations repetitive content and other 'junk' DNA can accumulate without placing the organism at a competitive disadvantage. There is little evidence to suggest that genome size is under strong widespread selection in multicellular eukaryotes. Genome size, independent of gene content, correlates poorly with most physiological traits and many eukaryotes, including mammals, harbor very large amounts of repetitive DNA.

However, birds likely have experienced strong selection for reduced genome size, in response to changing energetic needs for flight. Birds, unlike humans, produce nucleated red blood cells, and larger nuclei lead to lower levels of oxygen transport. Bird metabolism is far higher than that of mammals, due largely to flight, and oxygen needs are high. Hence, most birds have small, compact genomes with few repetitive elements. Indirect evidence suggests that non-avian theropod dinosaur ancestors of modern birds ^[3] also had reduced genome sizes, consistent with endothermy and high energetic needs for running speed. Many bacteria have also experienced selection for small genome size, as time of replication and energy consumption are so tightly correlated with fitness.

Repetitive elements

Transposable elements are self-replicating, selfish genetic elements which are capable of proliferating within host genomes. Many transposable elements are related to viruses, and share several proteins in common.

DNA transposons are cut and paste transposable elements which excise DNA and move it to alternate sections of the genome.

non-LTR retrotransposons

LTR retrotransposons

Helitrons

Alu elements comprise over XX % of the human genome. They are short non-autonomous repeat sequences.

Chromosome number and organization

The number of chromosomes in an organism's genome also does not necessarily correlate with the amount of DNA in its genome. The ant Myrmecia pilosula has only a single pair of chromosomes^[4] whereas the Adders-tongue fern Ophioglossum reticulatum has up to 1260 chromosomes.^[5] Cilliate genomes house each gene in individual chromosomes, resulting in a genome which is not physically linked. Reduced linkage through creation of additional chromosomes should effectively increase the efficiency of selection.

Changes in chromosome number can play a key role in speciation, as differing chromosome numbers can serve as a barrier to reproduction in hybrids. Human chromosome 2 was created from a fusion of two chimpanzee chromosomes and still contains central telomeres as well as a vestigial second centromere. Polyploidy especially allopolyploidy, which occurs often in plants, can also result in reproductive incompatibilities with parental species. Agrodiatus blue butterflies have diverse chromosome numbers ranging from n=10 to n=134 and additionally have one of the highest rates of speciation identified to date.^[6]

Gene content and distribution

Different organisms house different numbers of genes within their genomes as well as different patterns in the distribution of genes throughout the genome. Some organisms, such as most bacteria, Drosophila, and Arabidopsis have particularly compact genomes with little repetitive content or non-coding DNA. Other organisms, like mammals or maize, have large amounts of repetitive DNA, long introns, and substantial spacing between different genes. The content and distribution of genes within the genome can influence the rate at which certain types of mutations occur and can influence the subsequent evolution of different species. Genes with longer introns are more likely to recombine due to increased physical distance over the coding sequence. As such, long introns may facilitate ectopic recombination, and result in higher rates of new gene formation.

Organelles

In addition to the nuclear genome, endosymbiont organelles contain their own genetic material typically as circular plasmids. Mitochondrial and chloroplast DNA varies across taxa, but membrane-bound proteins, especially electron transport chain constituents are most often encoded in the organelle. Chloroplasts and mitochondria are maternally inherited in most species, as the organelles must pass through the egg. In a rare departure, some species of mussels are known to inherit mitochondria from father to son.

Origins of new genes

New genes arise from several different genetic mechanisms including gene duplication, de novo origination, retrotransposition, chimeric gene formation, recruitment of non-coding sequence, and gene truncation.

In gene duplication, a gene sequence is copied to create redundancy. Duplicated gene sequences can then mutate to develop new functions or to specialize so that each new gene performs a subset of the original ancestral functions. In addition to duplicating whole genes, sometimes only a domain or part of a protein is duplicated so that the resulting gene is an elongated version of the parental gene.

Retrotransposition creates new genes by copying mRNA to DNA and inserting it into the genome. Retrogenes often insert into new genomic locations, and often develop new expression patterns and functions.

Chimeric genes form when duplication, deletion, or incomplete retrotransposition combine portions of two different coding sequences to produce a novel gene sequence. Chimeras often cause regulatory changes and can shuffle protein domains to produce novel adaptive functions.

Novel genes can also arise from previously non-coding DNA.^[7] For instance, Levine and colleagues reported the origin of five new genes in the D. melanogaster genome from noncoding DNA.^[8]^[9] Similar de novo origin of genes has been also shown in other organisms such as yeast,^[10] rice^[11] and humans.^[12] De novo genes may evolve from transcripts that are already expressed at low levels.^[13] Mutation of a stop codon to a regular codon or a frameshift may cause an extended protein that includes a previously non-coding sequence.

Molecular phylogenetics

Molecular systematics is a product of the traditional field of systematics and molecular genetics. It uses DNA, RNA, or protein sequences to resolve questions in systematics, i.e. about their correct scientific classification or taxonomy from the point of view of evolutionary biology.
Molecular systematics has been made possible by the availability of techniques for DNA sequencing, which allow the determination of the exact sequence of nucleotides or bases in either DNA or RNA. At present it is still a long and expensive process to sequence the entire genome of an organism, and this has been done for only a few species. However, it is quite feasible to determine the sequence of a defined area of a particular chromosome. Typical molecular systematic analyses require the sequencing of around 1000 base pairs.

The driving forces of evolution

Depending on the relative importance assigned to the various forces of evolution, three perspectives provide evolutionary explanations for molecular evolution.^[14]
Selectionist hypotheses argue that selection is the driving force of molecular evolution. While acknowledging that many mutations are neutral, selectionists attribute changes in the frequencies of neutral alleles to linkage disequilibrium with other loci that are under selection, rather than to random genetic drift.^[15] Biases in codon usage are usually explained with reference to the ability of even weak selection to shape molecular evolution.^[16]

Neutralist hypotheses emphasize the importance of mutation, purifying selection and random genetic drift.^[17] The introduction of the neutral theory by Kimura,^[18] quickly followed by King and Jukes' own findings,^[19] led to a fierce debate about the relevance of neodarwinism at the molecular level. The Neutral theory of molecular evolution states that most mutations are deleterious and quickly removed by natural selection, but of the remaining ones, the vast majority are neutral with respect to fitness while the amount of advantageous mutations is vanishingly small. The fate of neutral mutations are governed by genetic drift, and contribute to both nucleotide polymorphism and fixed differences between species.^[20]^[21]^[22]

Mutationists hypotheses emphasize random drift and biases in mutation patterns.^[23] Sueoka was the first to propose a modern mutationist view. He proposed that the variation in GC content was not the result of positive selection, but a consequence of the GC mutational pressure.^[24]

Discordance with morphological evolution

There are sometimes discordances between molecular and morphological evolution, which are reflected in molecular and morphological systematic studies, especially of bacteria, archaea and eukaryotic microbes. These discordances can be categorized as two types: (i) one morphology, multiple lineages (e.g. morphological convergence, cryptic species) and (ii) one lineage, multiple morphologies (e.g. phenotypic plasticity, multiple life-cycle stages). Neutral evolution possibly could explain the incongruences in some cases.^[25]

Journals and societies

The Society for Molecular Biology and Evolution publishes the journals "Molecular Biology and Evolution" and "Genome Biology and Evolution" and holds an annual international meeting. Other journals dedicated to molecular evolution include Journal of Molecular Evolution and Molecular Phylogenetics and Evolution. Research in molecular evolution is also published in journals of genetics, molecular biology, genomics, systematics, and evolutionary biology.

Evolutionary psychology

From Wikipedia, the free encyclopedia

Evolutionary psychology (EP) is an approach in the social and natural sciences that examines psychological structure from a modern evolutionary perspective. It seeks to identify which human psychological traits are evolved adaptations – that is, the functional products of natural selection or sexual selection. Adaptationist thinking about physiological mechanisms, such as the heart, lungs, and immune system, is common in evolutionary biology. Some evolutionary psychologists apply the same thinking to psychology, arguing that the mind has a modular structure similar to that of the body, with different modular adaptations serving different functions. Evolutionary psychologists argue that much of human behavior is the output of psychological adaptations that evolved to solve recurrent problems in human ancestral environments.^[1]

Evolutionary psychologists suggest that EP is not simply a subdiscipline of psychology but that evolutionary theory can provide a foundational, metatheoretical framework that integrates the entire field of psychology, in the same way it has for biology.^[2]^[3]^[4]

Evolutionary psychologists hold that behaviors or traits that occur universally in all cultures are good candidates for evolutionary adaptations^[5] including the abilities to infer others' emotions, discern kin from non-kin, identify and prefer healthier mates, and cooperate with others. They report successful tests of theoretical predictions related to such topics as infanticide, intelligence, marriage patterns, promiscuity, perception of beauty, bride price, and parental investment.

The theories and findings of EP have applications in many fields, including economics, environment, health, law, management, psychiatry, politics, and literature.^[7]^[8]

Controversies concerning EP involve questions of testability, cognitive and evolutionary assumptions (such as modular functioning of the brain, and large uncertainty about the ancestral environment), importance of non-genetic and non-adaptive explanations, as well as political and ethical issues due to interpretations of research results.^[9]

Scope

Principles

Evolutionary psychology is an approach that views human nature as the product of a universal set of evolved psychological adaptations to recurring problems in the ancestral environment. Proponents of EP suggest that it seeks to integrate psychology into the other natural sciences, rooting it in the organizing theory of biology (evolutionary theory), and thus understanding psychology as a branch of biology. Anthropologist John Tooby and psychologist Leda Cosmides note:

Evolutionary psychology is the long-forestalled scientific attempt to assemble out of the disjointed, fragmentary, and mutually contradictory human disciplines a single, logically integrated research framework for the psychological, social, and behavioral sciences—a framework that not only incorporates the evolutionary sciences on a full and equal basis, but that systematically works out all of the revisions in existing belief and research practice that such a synthesis requires.^[10]

Just as human physiology and evolutionary physiology have worked to identify physical adaptations of the body that represent "human physiological nature," the purpose of evolutionary psychology is to identify evolved emotional and cognitive adaptations that represent "human psychological nature." According to Steven Pinker, EP is "not a single theory but a large set of hypotheses" and a term that "has also come to refer to a particular way of applying evolutionary theory to the mind, with an emphasis on adaptation, gene-level selection, and modularity."
Evolutionary psychology adopts an understanding of the mind that is based on the computational theory of mind. It describes mental processes as computational operations, so that, for example, a fear response is described as arising from a neurological computation that inputs the perceptional data, e.g. a visual image of a spider, and outputs the appropriate reaction, e.g. fear of possibly dangerous animals.

While philosophers have generally considered the human mind to include broad faculties, such as reason and lust, evolutionary psychologists describe evolved psychological mechanisms as narrowly focused to deal with specific issues, such as catching cheaters or choosing mates. EP views the human brain as comprising many functional mechanisms,^{[citation needed]} called psychological adaptations or evolved cognitive mechanisms or cognitive modules, designed by the process of natural selection. Examples include language-acquisition modules, incest-avoidance mechanisms, cheater-detection mechanisms, intelligence and sex-specific mating preferences, foraging mechanisms, alliance-tracking mechanisms, agent-detection mechanisms, and others. Some mechanisms, termed domain-specific, deal with recurrent adaptive problems over the course of human evolutionary history.^{[citation needed]} Domain-general mechanisms, on the other hand, are proposed to deal with evolutionary novelty.^{[citation needed]}

EP has roots in cognitive psychology and evolutionary biology but also draws on behavioral ecology, artificial intelligence, genetics, ethology, anthropology, archaeology, biology, and zoology. EP is closely linked to sociobiology,^[5] but there are key differences between them including the emphasis on domain-specific rather than domain-general mechanisms, the relevance of measures of current fitness, the importance of mismatch theory, and psychology rather than behavior. Most of what is now labeled as sociobiological research is now confined to the field of behavioral ecology.^{[citation needed]}

Nikolaas Tinbergen's four categories of questions can help to clarify the distinctions between several different, but complementary, types of explanations.^[11] Evolutionary psychology focuses primarily on the "why?" questions, while traditional psychology focuses on the "how?" questions.^[12]

		Sequential vs. Static Perspective
		Historical/Developmental Explanation of current form in terms of a historical sequence	Current Form Explanation of the current form of species
How vs. Why Questions	Proximate How an individual organism's structures function	Ontogeny Developmental explanations for changes in individuals, from DNA to their current form	Mechanism Mechanistic explanations for how an organism's structures work
How vs. Why Questions	Evolutionary Why a species evolved the structures (adaptations) it has	Phylogeny The history of the evolution of sequential changes in a species over many generations	Adaptation A species trait that evolved to solve a reproductive or survival problem in the ancestral environment

Premises

Evolutionary psychology is founded on several core premises.

The brain is an information processing device, and it produces behavior in response to external and internal inputs.^[2]^[13]
The brain's adaptive mechanisms were shaped by natural and sexual selection.^[2]^[13]
Different neural mechanisms are specialized for solving problems in humanity's evolutionary past.^[2]^[13]
The brain has evolved specialized neural mechanisms that were designed for solving problems that recurred over deep evolutionary time,^[13] giving modern humans stone-age minds.^[2]
Most contents and processes of the brain are unconscious; and most mental problems that seem easy to solve are actually extremely difficult problems that are solved unconsciously by complicated neural mechanisms.^[2]
Human psychology consists of many specialized mechanisms, each sensitive to different classes of information or inputs. These mechanisms combine to produce manifest behavior.^[13]

History

Nobel Laureates Nikolaas Tinbergen (left) and Konrad Lorenz (right) who were, with Karl von Frisch, acknowledged for work on animal behavior^[14]

Evolutionary psychology has its historical roots in Charles Darwin's theory of natural selection.^[5] In The Origin of Species, Darwin predicted that psychology would develop an evolutionary basis:

In the distant future I see open fields for far more important researches. Psychology will be based on a new foundation, that of the necessary acquirement of each mental power and capacity by gradation.

—Darwin, Charles (1859). The Origin of Species. Wikisource. p. 488.

Two of his later books were devoted to the study of animal emotions and psychology; The Descent of Man, and Selection in Relation to Sex in 1871 and The Expression of the Emotions in Man and Animals in 1872. Darwin's work inspired William James's functionalist approach to psychology.^[5] Darwin's theories of evolution, adaptation, and natural selection have provided insight into why brains function the way they do.^[15]^[16]

The content of EP has derived from, on one hand, the biological sciences (especially evolutionary theory as it relates to ancient human environments, the study of paleoanthropology and animal behavior) and, on the other, the human sciences, especially psychology.

Evolutionary biology as an academic discipline emerged with the modern evolutionary synthesis in the 1930s and 1940s.^[17] In the 1930s the study of animal behavior (ethology) emerged with the work of Dutch biologist Nikolaas Tinbergen and Austrian biologists Konrad Lorenz and Karl von Frisch.

W.D. Hamilton's (1964) papers on inclusive fitness and Robert Trivers's (1972)^[18] theories on reciprocity and parental investment helped to establish evolutionary thinking in psychology and the other social sciences. In 1975, Edward O. Wilson combined evolutionary theory with studies of animal and social behavior, building on the works of Lorenz and Tinbergen, in his book Sociobiology: The New Synthesis.

In the 1970s, two major branches developed from ethology. Firstly, the study of animal social behavior (including humans) generated sociobiology, defined by its pre-eminent proponent Edward O. Wilson in 1975 as "the systematic study of the biological basis of all social behavior"^[19] and in 1978 as "the extension of population biology and evolutionary theory to social organization."^[20] Secondly, there was behavioral ecology which placed less emphasis on social behavior by focusing on the ecological and evolutionary basis of both animal and human behavior.

In the 1970s and 1980s university departments began to include the term evolutionary biology in their titles. The modern era of evolutionary psychology was ushered in, in particular, by Donald Symons' 1979 book The Evolution of Human Sexuality and Leda Cosmides and John Tooby's 1992 book The Adapted Mind.^[5]

From psychology there are the primary streams of developmental, social and cognitive psychology. Establishing some measure of the relative influence of genetics and environment on behavior has been at the core of behavioral genetics and its variants, notably studies at the molecular level that examine the relationship between genes, neurotransmitters and behavior. Dual inheritance theory (DIT), developed in the late 1970s and early 1980s, has a slightly different perspective by trying to explain how human behavior is a product of two different and interacting evolutionary processes: genetic evolution and cultural evolution. DIT is seen by some as a "middle-ground" between views that emphasize human universals versus those that emphasize cultural variation.^[21]

Theoretical foundations

The theories on which evolutionary psychology is based originated with Charles Darwin's work, including his speculations about the evolutionary origins of social instincts in humans. Modern evolutionary psychology, however, is possible only because of advances in evolutionary theory in the 20th century.
Evolutionary psychologists say that natural selection has provided humans with many psychological adaptations, in much the same way that it generated humans' anatomical and physiological adaptations.^[22] As with adaptations in general, psychological adaptations are said to be specialized for the environment in which an organism evolved, the environment of evolutionary adaptedness, or EEA.^[22]^[23] Sexual selection provides organisms with adaptations related to mating.^[22] For male mammals, which have a relatively high maximal potential reproduction rate, sexual selection leads to adaptations that help them compete for females.^[22] For female mammals, with a relatively low maximal potential reproduction rate, sexual selection leads to choosiness, which helps females select higher quality mates.^[22] Charles Darwin described both natural selection and sexual selection, and he relied on group selection to explain the evolution of altruistic (self-sacrificing) behavior. But group selection was considered a weak explanation, because in any group the less altruistic individuals will be more likely to survive, and the group will become less self-sacrificing as a whole.

In 1964, William D. Hamilton proposed inclusive fitness theory, emphasizing a "gene's-eye" view of evolution. Hamilton noted that genes can increase the replication of copies of themselves into the next generation by influencing the organism's social traits in such a way that (statistically) results in helping the survival and reproduction of other copies of the same genes (most simply, identical copies in the organism's close relatives). According to "Hamilton's rule", self-sacrificing behaviors (and the genes influencing them) can evolve if they typically help the organism's close relatives so much that it more than compensates for the individual animal's sacrifice. Inclusive fitness theory resolved the issue of how "altruism" can evolve. Other theories also help explain the evolution of altruistic behavior, including evolutionary game theory, tit-for-tat reciprocity, and generalized reciprocity. These theories not only help explain the development of altruistic behavior, but also account for hostility toward cheaters (individuals that take advantage of others' altruism).^[24]

Several mid-level evolutionary theories inform evolutionary psychology. The r/K selection theory proposes that some species prosper by having many offspring, while others follow the strategy of having fewer offspring but investing much more in each one. Humans follow the second strategy. Parental investment theory explains how parents invest more or less in individual offspring based on how successful those offspring are likely to be, and thus how much they might improve the parents' inclusive fitness. According to the Trivers-Willard hypothesis, parents in good conditions tend to invest more in sons (who are best able to take advantage of good conditions), while parents in poor conditions tend to invest more in daughters (who are best able to have successful offspring even in poor conditions). According to life history theory, animals evolve life histories to match their environments, determining details such as age at first reproduction and number of offspring. Dual inheritance theory posits that genes and human culture have interacted, with genes affecting the development of culture, and culture, in turn, affecting human evolution on a genetic level (see also the Baldwin effect).

Evolved psychological mechanisms

Evolutionary psychology is based on the hypothesis that, just like hearts, lungs, livers, kidneys, and immune systems, cognition has functional structure that has a genetic basis, and therefore has evolved by natural selection. Like other organs and tissues, this functional structure should be universally shared amongst a species, and should solve important problems of survival and reproduction.
Evolutionary psychologists seek to understand psychological mechanisms by understanding the survival and reproductive functions they might have served over the course of evolutionary history.^{[citation needed]} These might include abilities to infer others' emotions, discern kin from non-kin, identify and prefer healthier mates, cooperate with others and follow leaders. Consistent with the theory of natural selection, evolutionary psychology sees humans as often in conflict with others, including mates and relatives. For instance, a mother may wish to wean her offspring from breastfeeding earlier than does her infant, which frees up the mother to invest in additional offspring.^[24]^[25] Evolutionary psychology also recognizes the role of kin selection and reciprocity in evolving prosocial traits such as altruism.^[24] Like chimps and bonobos, humans have subtle and flexible social instincts, allowing them to form extended families, lifelong friendships, and political alliances.^[24] In studies testing theoretical predictions, evolutionary psychologists have made modest findings on topics such as infanticide, intelligence, marriage patterns, promiscuity, perception of beauty, bride price and parental investment.

Products of evolution: adaptations, exaptations, byproducts, and random variation

Not all traits of organisms are adaptations. As noted in the table below, traits may also be exaptations, byproducts of adaptations (sometimes called "spandrels"), or random variation between individuals.^[26]

Psychological adaptations are hypothesized to be innate or relatively easy to learn, and to manifest in cultures worldwide. For example, the ability of toddlers to learn a language with virtually no training is likely to be a psychological adaptation. On the other hand, ancestral humans did not read or write, thus today, learning to read and write require extensive training, and presumably represent byproducts of cognitive processing that use psychological adaptations designed for other functions.^[27] However, variations in manifest behavior can result from universal mechanisms interacting with different local environments. For example, Caucasians who move from a northern climate to the equator will have darker skin. The mechanisms regulating their pigmentation do not change; rather the input to the those mechanisms change, resulting in different output.

	Adaptation	Exaptation	By-Product	Random Variation
Definition	Organismic trait designed to solve an ancestral problem(s). Shows complexity, special "design", functionality	Adaptation that has been "re-purposed" to solve a different adaptive problem.	Byproduct of an adaptive mechanism with no current or ancestral function	Random variations in an adaptation or byproduct
Physiological Example	Bones / Umbilical cord	Small bones of the inner ear	White color of bones / Belly button	Bumps on the skull, convex or concave belly button shape
Psychological Example	Toddlers’ ability to learn to talk with minimal instruction.	Voluntary Attention	Ability to learn to read and write.	Within-sex variations in voice pitch.

One of the tasks of evolutionary psychology is to identify which psychological traits are likely to be adaptations, byproducts or random variation. George C Williams suggested that an "adaptation is a special and onerous concept that should only be used where it is really necessary."^[28] As noted by Williams and others, adaptations can be identified by their improbable complexity, species universality, and adaptive functionality.

Obligate and facultative adaptations

A question that may be asked about an adaptation is whether it is generally obligate (relatively robust in the face of typical environmental variation) or facultative (sensitive to typical environmental variation).^[29] The sweet taste of sugar and the pain of hitting one's knee against concrete are the result of fairly obligate psychological adaptations; typical environmental variability during development does not much affect their operation. By contrast, facultative adaptations are somewhat like "if-then" statements. For example, adult attachment style seems particularly sensitive to early childhood experiences. As adults, the propensity to develop close, trusting bonds with others is dependent on whether early childhood caregivers could be trusted to provide reliable assistance and attention. The adaptation for skin to tan is conditional to exposure to sunlight; this is an example of another facultative adaptation. When a psychological adaptation is facultative, evolutionary psychologists concern themselves with how developmental and environmental inputs influence the expression of the adaptation.

Cultural universals

Evolutionary psychologists hold that behaviors or traits that occur universally in all cultures are good candidates for evolutionary adaptations.^[5] Cultural universals include behaviors related to language, cognition, social roles, gender roles, and technology.^[30] Evolved psychological adaptations (such as the ability to learn a language) interact with cultural inputs to produce specific behaviors (e.g., the specific language learned). Basic gender differences, such as greater eagerness for sex among men and greater coyness among women,^[31] are explained as sexually dimorphic psychological adaptations that reflect the different reproductive strategies of males and females.^[24]^[32] Evolutionary psychologists contrast their approach to what they term the "standard social science model," according to which the mind is a general-purpose cognition device shaped almost entirely by culture.^[33]^[34]

Environment of evolutionary adaptedness

EP argues that to properly understand the functions of the brain, one must understand the properties of the environment in which the brain evolved. That environment is often referred to as the "environment of evolutionary adaptedness" (EEA).^[23]
The idea of an environment of evolutionary adaptedness was first explored as a part of attachment theory by John Bowlby.^[35] This is the environment to which a particular evolved mechanism is adapted. More specifically, the EEA is defined as the set of historically recurring selection pressures that formed a given adaptation, as well as those aspects of the environment that were necessary for the proper development and functioning of the adaptation.

Humans, comprising the genus Homo, appeared between 1.5 and 2.5 million years ago, a time that roughly coincides with the start of the Pleistocene 2.6 million years ago. Because the Pleistocene ended a mere 12,000 years ago, most human adaptations either newly evolved during the Pleistocene, or were maintained by stabilizing selection during the Pleistocene. Evolutionary psychology therefore proposes that the majority of human psychological mechanisms are adapted to reproductive problems frequently encountered in Pleistocene environments.^[36] In broad terms, these problems include those of growth, development, differentiation, maintenance, mating, parenting, and social relationships.

The EEA is significantly different from modern society.^[37] The ancestors of modern humans lived in smaller groups, had more cohesive cultures, and had more stable and rich contexts for identity and meaning.^[37] Researchers look to existing hunter-gatherer societies for clues as to how hunter-gatherers lived in the EEA.^[24] Unfortunately, the few surviving hunter-gatherer societies are different from each other, and they have been pushed out of the best land and into harsh environments, so it is not clear how closely they reflect ancestral culture.^[24]

Evolutionary psychologists sometimes look to chimpanzees, bonobos, and other great apes for insight into human ancestral behavior.^[24] Christopher Ryan and Cacilda Jetha argue that evolutionary psychologists have overemphasized the similarity of humans and chimps, which are more violent, while underestimating the similarity of humans and bonobos, which are more peaceful.^[38]

Mismatches

Since an organism's adaptations were suited to its ancestral environment, a new and different environment can create a mismatch. Because humans are mostly adapted to Pleistocene environments, psychological mechanisms sometimes exhibit "mismatches" to the modern environment. One example is the fact that although about 10,000 people are killed with guns in the US annually,^[39] whereas spiders and snakes kill only a handful, people nonetheless learn to fear spiders and snakes about as easily as they do a pointed gun, and more easily than an unpointed gun, rabbits or flowers.^[40] A potential explanation is that spiders and snakes were a threat to human ancestors throughout the Pleistocene, whereas guns (and rabbits and flowers) were not. There is thus a mismatch between humans' evolved fear-learning psychology and the modern environment.^[41]^[42]

This mismatch also shows up in the phenomena of the supernormal stimulus, a stimulus that elicits a response more strongly than the stimulus for which the response evolved. The term was coined by Niko Tinbergen to refer to non-human animal behavior, but psychologist Deirdre Barrett said that supernormal stimulation governs the behavior of humans as powerfully as that of other animals. She explained junk food as an exaggerated stimulus to cravings for salt, sugar, and fats,^[43] and she says that television is an exaggeration of social cues of laughter, smiling faces and attention-grabbing action.^[44] Magazine centerfolds and double cheeseburgers pull instincts intended for an EEA where breast development was a sign of health, youth and fertility in a prospective mate, and fat was a rare and vital nutrient.^[45] Psychologist Mark van Vugt recently argued that modern organizational leadership is a mismatch.^[46] His argument is that humans are not adapted to work in large, anonymous bureaucratic structures with formal hierarchies. The human mind still responds to personalized, charismatic leadership primarily in the context of informal, egalitarian settings. Hence the dissatisfaction and alienation that many employees experience. Salaries, bonuses and other privileges exploit instincts for relative status, which attract particularly males to senior executive positions.^[47]

Research methods

Evolutionary theory is heuristic in that it may generate hypotheses that might not be developed from other theoretical approaches. One of the major goals of adaptationist research is to identify which organismic traits are likely to be adaptations, and which are byproducts or random variations. As noted earlier, adaptations are expected to show evidence of complexity, functionality, and species universality, while byproducts or random variation will not. In addition, adaptations are expected to manifest as proximate mechanisms that interact with the environment in either a generally obligate or facultative fashion (see above). Evolutionary psychologists are also interested in identifying these proximate mechanisms (sometimes termed "mental mechanisms" or "psychological adaptations") and what type of information they take as input, how they process that information, and their outputs.^[29]
Evolutionary developmental psychology, or "evo-devo," focuses on how adaptations may be activated at certain developmental times (e.g., losing baby teeth, adolescence, etc.) or how events during the development of an individual may alter life history trajectories.

Evolutionary psychologists use several strategies to develop and test hypotheses about whether a psychological trait is likely to be an evolved adaptation. Buss (2011)^[48] notes that these methods include:

Cross-cultural Consistency. Characteristics that have been demonstrated to be cross cultural human universals such as smiling, crying, facial expressions are presumed to be evolved psychological adaptations. Several evolutionary psychologists have collected massive datasets from cultures around the world to assess cross-cultural universality.

Function to Form (or "problem to solution"). The fact that males, but not females, risk potential misidentification of genetic offspring (referred to as "paternity insecurity") led evolutionary psychologists to hypothesize that, compared to females, male jealousy would be more focused on sexual, rather than emotional, infidelity.

Form to Function (reverse-engineering – or "solution to problem"). Morning sickness, and associated aversions to certain types of food, during pregnancy seemed to have the characteristics of an evolved adaptation (complexity and universality). Margie Profet hypothesized that the function was to avoid the ingestion of toxins during early pregnancy that could damage fetus (but which are otherwise likely to be harmless to healthy non-pregnant women).

Corresponding Neurological Modules. Evolutionary psychology and cognitive neuropsychology are mutually compatible – evolutionary psychology helps to identify psychological adaptations and their ultimate, evolutionary functions, while neuropsychology helps to identify the proximate manifestations of these adaptations.

Evolutionary psychologists also use various sources of data for testing, including experiments, archaeological records, data from hunter-gatherer societies, observational studies, neuroscience data, self-reports and surveys, public records, and human products.^[49] Recently, additional methods and tools have been introduced based on fictional scenarios,^[50] mathematical models,^[51] and multi-agent computer simulations.^[52]

Major areas of research

Foundational areas of research in evolutionary psychology can be divided into broad categories of adaptive problems that arise from the theory of evolution itself: survival, mating, parenting, family and kinship, interactions with non-kin, and cultural evolution.

Survival and individual level psychological adaptations

Problems of survival are thus clear targets for the evolution of physical and psychological adaptations.^{[clarification needed]} Major problems the ancestors of present day humans faced included food selection and acquisition; territory selection and physical shelter; and avoiding predators and other environmental threats.^[29]

Consciousness

Consciousness meets George Williams' criteria of species universality, complexity,^[53] and functionality, and it is a trait that apparently increases fitness.^[54]
In his paper "Evolution of consciousness," John Eccles argues that special anatomical and physical adaptations of the mammalian cerebral cortex gave rise to consciousness.^[55] In contrast, others have argued that the recursive circuitry underwriting consciousness is much more primitive, having evolved initially in pre-mammalian species because it improves the capacity for interaction with both social and natural environments by providing an energy-saving "neutral" gear in an otherwise energy-expensive motor output machine.^[56] Once in place, this recursive circuitry may well have provided a basis for the subsequent development of many of the functions that consciousness facilitates in higher organisms, as outlined by Bernard J. Baars.^[57] Richard Dawkins suggested that humans evolved consciousness in order to make themselves the subjects of thought.^[58] Daniel Povinelli suggests that large, tree-climbing apes evolved consciousness to take into account one's own mass when moving safely among tree branches.^[58] Consistent with this hypothesis, Gordon Gallup found that chimps and orangutans, but not little monkeys or terrestrial gorillas, demonstrated self-awareness in mirror tests.^[58]

The concept of consciousness can refer to voluntary action, awareness, or wakefulness. However, even voluntary behavior involves unconscious mechanisms. Many cognitive processes take place in the cognitive unconscious, unavailable to conscious awareness. Some behaviors are conscious when learned but then become unconscious, seemingly automatic. Learning, especially implicitly learning a skill, can take place outside of consciousness. For example, plenty of people know how to turn right when they ride a bike, but very few can accurately explain how they actually do so. Evolutionary psychology approaches self-deception as an adaptation that can improve one's results in social exchanges.^[58]

Sleep may have evolved to conserve energy when activity would be less fruitful or more dangerous, such as at night, especially in winter.^[58]

Sensation and perception

Many experts, such as Jerry Fodor, write that the purpose of perception is knowledge, but evolutionary psychologists hold that its primary purpose is to guide action.^[59] For example, they say, depth perception seems to have evolved not to help us know the distances to other objects but rather to help us move around in space.^[59]
Evolutionary psychologists say that animals from fiddler crabs to humans use eyesight for collision avoidance, suggesting that vision is basically for directing action, not providing knowledge.^[59]

Building and maintaining sense organs is metabolically expensive, so these organs evolve only when they improve an organism's fitness.^[59] More than half the brain is devoted to processing sensory information, and the brain itself consumes roughly one-fourth of one's metabolic resources, so the senses must provide exceptional benefits to fitness.^[59] Perception accurately mirrors the world; animals get useful, accurate information through their senses.^[59]

Scientists who study perception and sensation have long understood the human senses as adaptations.^[59] Depth perception consists of processing over half a dozen visual cues, each of which is based on a regularity of the physical world.^[59] Vision evolved to respond to the narrow range of electromagnetic energy that is plentiful and that does not pass through objects.^[59] Sound waves go around corners and interact with obstacles, creating a complex pattern that includes useful information about the sources of and distances to objects.^[59] Larger animals naturally make lower-pitched sounds as a consequence of their size.^[59] The range over which an animal hears, on the other hand, is determined by adaptation. Homing pigeons, for example, can hear very low-pitched sound (infrasound) that carries great distances, even though most smaller animals detect higher-pitched sounds.^[59] Taste and smell respond to chemicals in the environment that are thought to have been significant for fitness in the EEA.^[59] For example, salt and sugar were apparently both valuable to the human or pre-human inhabitants of the EEA, so present day humans have an intrinsic hunger for salty and sweet tastes.^[59] The sense of touch is actually many senses, including pressure, heat, cold, tickle, and pain.^[59] Pain, while unpleasant, is adaptive.^[59] An important adaptation for senses is range shifting, by which the organism becomes temporarily more or less sensitive to sensation.^[59] For example, one's eyes automatically adjust to dim or bright ambient light.^[59] Sensory abilities of different organisms often coevolve, as is the case with the hearing of echolocating bats and that of the moths that have evolved to respond to the sounds that the bats make.^[59]

Evolutionary psychologists contend that perception demonstrates the principle of modularity, with specialized mechanisms handling particular perception tasks.^[59] For example, people with damage to a particular part of the brain suffer from the specific defect of not being able to recognize faces (prosopagnosia).^[59] EP suggests that this indicates a so-called face-reading module.^[59]

Learning and facultative adaptations

In evolutionary psychology, learning is said to be accomplished through evolved capacities, specifically facultative adaptations.^[60] Facultative adaptations express themselves differently depending on input from the environment.^[60] Sometimes the input comes during development and helps shape that development.^[60] For example, migrating birds learn to orient themselves by the stars during a critical period in their maturation.^[60] Evolutionary psychologists believe that humans also learn language along an evolved program, also with critical periods.^[60] The input can also come during daily tasks, helping the organism cope with changing environmental conditions.^[60] For example, animals evolved Pavlovian conditioning in order to solve problems about causal relationships.^[60] Animals accomplish learning tasks most easily when those tasks resemble problems that they faced in their evolutionary past, such as a rat learning where to find food or water.^[60] Learning capacities sometimes demonstrate differences between the sexes.^[60] In many animal species, for example, males can solve spatial problem faster and more accurately than females, due to the effects of male hormones during development.^[60] The same might be true of humans.^[60]

Emotion and motivation

Motivations direct and energize behavior, while emotions provide the affective component to motivation, positive or negative.^[61] In the early 1970s, Paul Ekman and colleagues began a line of research which suggests that many emotions are universal.^[61] He found evidence that humans share at least five basic emotions: fear, sadness, happiness, anger, and disgust.^[61] Social emotions evidently evolved to motivate social behaviors that were adaptive in the EEA.^[61] For example, spite seems to work against the individual but it can establish an individual's reputation as someone to be feared.^[61] Shame and pride can motivate behaviors that help one maintain one's standing in a community, and self-esteem is one's estimate of one's status.^[24]^[61] Motivation has a neurobiologial basis in the reward system of the brain. Recently, it has been suggested that reward systems may evolve in such a way that there may be an inherent or unavoidable trade-off in the motivational system for activities of short versus long duration.^[62]

Cognition

Cognition refers to internal representations of the world and internal information processing. From an EP perspective, cognition is not "general purpose," but uses heuristics, or strategies, that generally increase the likelihood of solving problems that the ancestors of present day humans routinely faced. For example, present day humans are far more likely to solve logic problems that involve detecting cheating (a common problem given humans' social nature) than the same logic problem put in purely abstract terms.^[63] Since the ancestors of present day humans did not encounter truly random events, present day humans may be cognitively predisposed to incorrectly identify patterns in random sequences. "Gamblers' Fallacy" is one example of this. Gamblers may falsely believe that they have hit a "lucky streak" even when each outcome is actually random and independent of previous trials. Most people believe that if a fair coin has been flipped 9 times and Heads appears each time, that on the tenth flip, there is a greater than 50% chance of getting Tails.^[61] Humans find it far easier to make diagnoses or predictions using frequency data than when the same information is presented as probabilities or percentages, presumably because the ancestors of present day humans lived in relatively small tribes (usually with fewer than 150 people) where frequency information was more readily available.^[61]

Personality

Evolutionary psychology is primarily interested in finding commonalities between people, or basic human psychological nature. From an evolutionary perspective, the fact that people have fundamental differences in personality traits initially presents something of a puzzle.^[64] (Note: The field of behavioral genetics is concerned with statistically partitioning differences between people into genetic and environmental sources of variance. However, understanding the concept of heritability can be tricky—heritability refers only to the differences between people, never the degree to which the traits of an individual are due to environmental or genetic factors, since traits are always a complex interweaving of both.)

Personality traits are conceptualized by evolutionary psychologists as due to normal variation around an optimum, due to frequency-dependent selection (behavioral polymorphisms), or as facultative adaptations. Like variability in height, some personality traits may simply reflect inter-individual variability around a general optimum.^[64] Or, personality traits may represent different genetically predisposed "behavioral morphs" – alternate behavioral strategies that depend on the frequency of competing behavioral strategies in the population. For example, if most of the population is generally trusting and gullible, the behavioral morph of being a "cheater" (or, in the extreme case, a sociopath) may be advantageous.^[65] Finally, like many other psychological adaptations, personality traits may be facultative—sensitive to typical variations in the social environment, especially during early development. For example, later born children are more likely than first borns to be rebellious, less conscientious and more open to new experiences, which may be advantageous to them given their particular niche in family structure.^[66] It is important to note that shared environmental influences do play a role in personality and are not always of less importance than genetic factors. However, shared environmental influences often decrease to near zero after adolescence but do not completely disappear.^[67]

Language

According to Steven Pinker, who builds on the work by Noam Chomsky, the universal human ability to learn to talk between the ages of 1 – 4, basically without training, suggests that language acquisition is a distinctly human psychological adaptation (see, in particular, Pinker's The Language Instinct). Pinker and Bloom (1990) argue that language as a mental faculty shares many likenesses with the complex organs of the body which suggests that, like these organs, language has evolved as an adaptation, since this is the only known mechanism by which such complex organs can develop.^[68]
Pinker follows Chomsky in arguing that the fact that children can learn any human language with no explicit instruction suggests that language, including most of grammar, is basically innate and that it only needs to be activated by interaction. Chomsky himself does not believe language to have evolved as an adaptation, but suggests that it likely evolved as a byproduct of some other adaptation, a so-called spandrel. But Pinker and Bloom argue that the organic nature of language strongly suggests that it has an adaptational origin.^[69]

Evolutionary psychologists hold that the FOXP2 gene may well be associated with the evolution of human language.^[70] In the 1980s, psycholinguist Myrna Gropnik identified a dominant gene that causes language impairment in the KE family of Britain.^[70] This gene turned out to be a mutation of the FOXP2 gene.^[70] Humans have a unique allele of this gene, which has otherwise been closely conserved through most of mammalian evolutionary history.^[70] This unique allele seems to have first appeared between 100 and 200 thousand years ago, and it is now all but universal in humans.^[70] However, the once-popular idea that FOXP2 is a 'grammar gene' or that it triggered the emergence of language in Homo sapiens is now widely discredited.^[71]

Currently several competing theories about the evolutionary origin of language coexist, none of them having achieved a general consensus.^[72] Researchers of language acquisition in primates and humans such as Michael Tomasello and Talmy Givón, argue that the innatist framework has understated the role of imitation in learning and that it is not at all necessary to posit the existence of an innate grammar module to explain human language acquisition. Tomasello argues that studies of how children and primates actually acquire communicative skills suggests that humans learn complex behavior through experience, so that instead of a module specifically dedicated to language acquisition, language is acquired by the same cognitive mechanisms that are used to acquire all other kinds of socially transmitted behavior.^[73]

On the issue of whether language is best seen as having evolved as an adaptation or as a spandrel, evolutionary biologist W. Tecumseh Fitch, following Stephen J. Gould, argues that it is unwarranted to assume that every aspect of language is an adaptation, or that language as a whole is an adaptation. He criticizes some strands of evolutionary psychology for suggesting a pan-adaptionist view of evolution, and dismisses Pinker and Bloom's question of whether "Language has evolved as an adaptation" as being misleading. He argues instead that from a biological viewpoint the evolutionary origins of language is best conceptualized as being the probable result of a convergence of many separate adaptations into a complex system.^[74] A similar argument is made by Terrence Deacon who in The Symbolic Species argues that the different features of language have co-evolved with the evolution of the mind and that the ability to use symbolic communication is integrated in all other cognitive processes.^[75]

If the theory that language could have evolved as a single adaptation is accepted, the question becomes which of its many functions has been the basis of adaptation, several evolutionary hypotheses have been posited: that it evolved for the purpose of social grooming, that it evolved to as a way to show mating potential or that it evolved to form social contracts. Evolutionary psychologists recognize that these theories are all speculative and that much more evidence is required to understand how language might have been selectively adapted.^[76]

Mating

Given that sexual reproduction is the means by which genes are propagated into future generations, sexual selection plays a large role in human evolution. Human mating, then, is of interest to evolutionary psychologists who aim to investigate evolved mechanisms to attract and secure mates.^[77] Several lines of research have stemmed from this interest, such as studies of mate selection^[78]^[79]^[80] mate poaching,^[81] mate retention,^[82] mating preferences^[83] and conflict between the sexes.^[84]
In 1972 Robert Trivers published an influential paper^[85] on sex differences that is now referred to as parental investment theory. The size differences of gametes (anisogamy) is the fundamental, defining difference between males (small gametes—sperm) and females (large gametes—ova). Trivers noted that anisogamy typically results in different levels of parental investment between the sexes, with females initially investing more. Trivers proposed that this difference in parental investment leads to the sexual selection of different reproductive strategies between the sexes and to sexual conflict. For example, he suggested that the sex that invests less in offspring will generally compete for access to the higher-investing sex to increase their inclusive fitness (also see Bateman's principle^[86]). Trivers posited that differential parental investment led to the evolution sexual dimorphisms in mate choice, intra- and inter- sexual reproductive competition, and courtship displays. In mammals, including humans, females make a much larger parental investment than males (i.e. gestation followed by childbirth and lactation). Parental investment theory is a branch of life history theory.

Buss and Schmitt's (1993) Sexual Strategies Theory ^[87] proposed that, due to differential parental investment, humans have evolved sexually dimorphic adaptations related to "sexual accessibility, fertility assessment, commitment seeking and avoidance, immediate and enduring resource procurement, paternity certainty, assessment of mate value, and parental investment." Their Strategic Interference Theory^[88] suggested that conflict between the sexes occurs when the preferred reproductive strategies of one sex interfere with those of the other sex, resulting in the activation of emotional responses such as anger or jealousy.

Women are generally more selective when choosing mates, especially under short-term mating conditions. However, under some circumstances, short term mating can provide benefits to women as well, such as fertility insurance, trading up to better genes, reducing risk of inbreeding, and insurance protection of her offspring.^[89]

Due to male paternity insecurity, sex differences have been found in such domains as sexual jealousy.^[90]^[91] Females generally react more adversely to emotional infidelity and males will react more to sexual infidelity. This particular pattern is predicted because the costs involved in mating for each sex are distinct. Women, on average, should prefer a mate who can offer resources (e.g., financial, commitment), thus, a woman risks losing such resources with a mate who commits emotional infidelity. Men, on the other hand, are never certain of the genetic paternity of their children because they do not bear the offspring themselves ("paternity insecurity"). This suggests that for men sexual infidelity would generally be more aversive than emotional infidelity because investing resources in another man's offspring does not lead to propagation of their own genes.^[92]

Another interesting line of research is that which examines women's mate preferences across the ovulatory cycle.^[93]^[94] The theoretical underpinning of this research is that ancestral women would have evolved mechanisms to select mates with certain traits depending on their hormonal status. For example, the theory hypothesizes that, during the ovulatory phase of a woman's cycle (approximately days 10–15 of a woman's cycle),^[95] a woman who mated with a male with high genetic quality would have been more likely, on average, to produce and rear a healthy offspring than a woman who mated with a male with low genetic quality. These putative preferences are predicted to be especially apparent for short-term mating domains because a potential male mate would only be offering genes to a potential offspring. This hypothesis allows researchers to examine whether women select mates who have characteristics that indicate high genetic quality during the high fertility phase of their ovulatory cycles. Indeed, studies have shown that women's preferences vary across the ovulatory cycle. In particular, Haselton and Miller (2006) showed that highly fertile women prefer creative but poor men as short-term mates. Creativity may be a proxy for good genes.^[96] Research by Gangestad et al. (2004) indicates that highly fertile women prefer men who display social presence and intrasexual competition; these traits may act as cues that would help women predict which men may have, or would be able to acquire, resources.

Parenting

Reproduction is always costly for women, and can also be for men. Individuals are limited in the degree to which they can devote time and resources to producing and raising their young, and such expenditure may also be detrimental to their future condition, survival and further reproductive output. Parental investment is any parental expenditure (time, energy etc.) that benefits one offspring at a cost to parents' ability to invest in other components of fitness (Clutton-Brock 1991: 9; Trivers 1972). Components of fitness (Beatty 1992) include the well being of existing offspring, parents' future reproduction, and inclusive fitness through aid to kin (Hamilton, 1964). Parental investment theory is a branch of life history theory.
Robert Trivers' theory of parental investment predicts that the sex making the largest investment in lactation, nurturing and protecting offspring will be more discriminating in mating and that the sex that invests less in offspring will compete for access to the higher investing sex (see Bateman's principle).^[86] Sex differences in parental effort are important in determining the strength of sexual selection.

The benefits of parental investment to the offspring are large and are associated with the effects on condition, growth, survival and ultimately, on reproductive success of the offspring. However, these benefits can come at the cost of parent's ability to reproduce in the future e.g. through the increased risk of injury when defending offspring against predators, the loss of mating opportunities whilst rearing offspring and an increase in the time to the next reproduction. Overall, parents are selected to maximize the difference between the benefits and the costs, and parental care will be likely to evolve when the benefits exceed the costs.

The Cinderella effect is an alleged high incidence of stepchildren being physically, emotionally or sexually abused, neglected, murdered, or otherwise mistreated at the hands of their stepparents at significantly higher rates than their genetic counterparts. It takes its name from the fairy tale character Cinderella, who in the story was cruelly mistreated by her stepmother and stepsisters.^[97] Daly and Wilson (1996) noted: "Evolutionary thinking led to the discovery of the most important risk factor for child homicide – the presence of a stepparent. Parental efforts and investments are valuable resources, and selection favors those parental psyches that allocate effort effectively to promote fitness. The adaptive problems that challenge parental decision making include both the accurate identification of one's offspring and the allocation of one's resources among them with sensitivity to their needs and abilities to convert parental investment into fitness increments…. Stepchildren were seldom or never so valuable to one's expected fitness as one's own offspring would be, and those parental psyches that were easily parasitized by just any appealing youngster must always have incurred a selective disadvantage"(Daly & Wilson, 1996, pp. 64–65). However, they note that not all stepparents will "want" to abuse their partner's children, or that genetic parenthood is any insurance against abuse. They see step parental care as primarily "mating effort" towards the genetic parent.^[98]

Family and kin

Inclusive fitness is the sum of an organism's classical fitness (how many of its own offspring it produces and supports) and the number of equivalents of its own offspring it can add to the population by supporting others.^[99] The first component is called classical fitness by Hamilton (1964).

From the gene's point of view, evolutionary success ultimately depends on leaving behind the maximum number of copies of itself in the population. Until 1964, it was generally believed that genes only achieved this by causing the individual to leave the maximum number of viable offspring. However, in 1964 W. D. Hamilton proved mathematically that, because close relatives of an organism share some identical genes, a gene can also increase its evolutionary success by promoting the reproduction and survival of these related or otherwise similar individuals.

Hamilton concluded that this leads natural selection to favor organisms that would behave in ways that maximize their inclusive fitness. It is also true that natural selection favors behavior that maximizes personal fitness.
Hamilton's rule describes mathematically whether or not a gene for altruistic behavior will spread in a population:

rb > c \

where

$c \$ is the reproductive cost to the altruist,
$b \$ is the reproductive benefit to the recipient of the altruistic behavior, and
$r \$ is the probability, above the population average, of the individuals sharing an altruistic gene – commonly viewed as "degree of relatedness".

The concept serves to explain how natural selection can perpetuate altruism. If there is an '"altruism gene"' (or complex of genes) that influences an organism's behavior to be helpful and protective of relatives and their offspring, this behavior also increases the proportion of the altruism gene in the population, because relatives are likely to share genes with the altruist due to common descent. Altruists may also have some way to recognize altruistic behavior in unrelated individuals and be inclined to support them. As Dawkins points out in The Selfish Gene (Chapter 6) and The Extended Phenotype,^[100] this must be distinguished from the green-beard effect.
Although it is generally true that humans tend to be more altruistic toward their kin than toward non-kin, the relevant proximate mechanisms that mediate this cooperation have been debated (see kin recognition), with some arguing that kin status is determined primarily via social and cultural factors (such as co-residence, maternal association of sibs, etc.),^[101] while others have argued that kin recognition can also mediated by biological factors such as facial resemblance and immunogenetic similarity of the major histocompatibility complex (MHC).^[102] For a discussion of the interaction of these social and biological kin recognition factors see Lieberman, Tooby, and Cosmides (2007)^[103] (PDF).

Whatever the proximate mechanisms of kin recognition there is substantial evidence that humans act generally more altruistically to close genetic kin compared to genetic non-kin.^[104]^[105]^[106]

Interactions with non-kin / reciprocity

Although interactions with non-kin are generally less altruistic compared to those with kin, cooperation can be maintained with non-kin via mutually beneficial reciprocity as was proposed by Robert Trivers.^[18] If there are repeated encounters between the same two players in an evolutionary game in which each of them can choose either to "cooperate" or "defect," then a strategy of mutual cooperation may be favored even if it pays each player, in the short term, to defect when the other cooperates. Direct reciprocity can lead to the evolution of cooperation only if the probability, w, of another encounter between the same two individuals exceeds the cost-to-benefit ratio of the altruistic act:

w > c/b

Reciprocity can also be indirect if information about previous interactions is shared. Reputation allows evolution of cooperation by indirect reciprocity. Natural selection favors strategies that base the decision to help on the reputation of the recipient: studies show that people who are more helpful are more likely to receive help. The calculations of indirect reciprocity are complicated and only a tiny fraction of this universe has been uncovered, but again a simple rule has emerged.^[107] Indirect reciprocity can only promote cooperation if the probability, q, of knowing someone’s reputation exceeds the cost-to-benefit ratio of the altruistic act:

q > c/b

One important problem with this explanation is that individuals may be able to evolve the capacity to obscure their reputation, reducing the probability, q, that it will be known.^[108]

Trivers argues that friendship and various social emotions evolved in order to manage reciprocity.^[109] Liking and disliking, he says, evolved to help present day humans' ancestors form coalitions with others who reciprocated and to exclude those who did not reciprocate.^[109] Moral indignation may have evolved to prevent one's altruism from being exploited by cheaters, and gratitude may have motivated present day humans' ancestors to reciprocate appropriately after benefiting from others' altruism.^[109] Likewise, present day humans feel guilty when they fail to reciprocate.^[109] These social motivations match what evolutionary psychologists expect to see in adaptations that evolved to maximize the benefits and minimize the drawbacks of reciprocity.^[109]

Evolutionary psychologists say that humans have psychological adaptations that evolved specifically to help us identify nonreciprocators, commonly referred to as "cheaters."^[109] In 1993, Robert Frank and his associates found that participants in a prisoner's dilemma scenario were often able to predict whether their partners would "cheat," based on a half hour of unstructured social interaction.^[109] In a 1996 experiment, for example, Linda Mealey and her colleagues found that people were better at remembering the faces of people when those faces were associated with stories about those individuals cheating (such as embezzling money from a church).^[109]

Strong reciprocity (or "tribal reciprocity")

Humans may have an evolved set of psychological adaptations that predispose them to be more cooperative than otherwise would be expected with members of their tribal in-group, and, more nasty to members of tribal out groups. These adaptations may have be a consequent of tribal warfare.^[110] Humans may also have predispositions for "altruistic punishment"—to punish in-group members who violate in-group rules, even when this altruistic behavior cannot be justified in terms of helping those you are related to (kin selection), cooperating with those who you will interact with again (direct reciprocity), or cooperating to better your reputation with others (indirect reciprocity).^[111]^[112]

Evolution and culture

Memetics is a theory of mental content based on an analogy with evolution, originating from Richard Dawkins' 1976 book The Selfish Gene. It purports to be an approach to evolutionary models of cultural information transfer. A meme, analogous to a gene, is essentially a "unit of culture"—an idea, belief, pattern of behavior, etc. which is "hosted" in one or more individual minds, and which can reproduce itself from mind to mind. Thus what would otherwise be regarded as one individual influencing another to adopt a belief is seen memetically as a meme reproducing itself. As with genetics, particularly under Dawkins's interpretation, a meme's success may be due to its contribution to the effectiveness of its host. Memetics is notable for sidestepping the traditional concern with the truth of ideas and beliefs.
Susan Blackmore (2002) re-stated the definition of meme as: whatever is copied from one person to another person, whether habits, skills, songs, stories, or any other kind of information. Further she said that memes, like genes, are replicators in the sense as defined by Dawkins.^[113] That is, they are information that is copied. Memes are copied by imitation, teaching and other methods. The copies are not perfect: memes are copied with variation; moreover, memes compete for humans' limited memory capacity and for the chance to be copied again. Only some of the variants can survive. The combination of these three elements (copies; variation; competition for survival) forms precisely the condition for Darwinian evolution, and so memes (and hence human cultures) evolve. Large groups of memes that are copied and passed on together are called co-adapted meme complexes, or memeplexes. In her definition, the way that a meme replicates is through imitation.

Dual inheritance theory (DIT), also known as gene-culture coevolution, suggests that cultural information and genes co-evolve. Marcus Feldman and Luigi Luca Cavalli-Sforza (1976) published perhaps the first dynamic models of gene-culture coevolution.^[114] These models were to form the basis for subsequent work on DIT, heralded by the publication of three seminal books in 1980 and 1981. Charles Lumsden and E.O. Wilson's Genes, Mind and Culture (1981).^[115] also outlined a series of mathematical models of how genetic evolution might favor the selection of cultural traits and how cultural traits might, in turn, affect the speed of genetic evolution. Another 1981 book relevant to this topic was Cavalli-Sforza and Feldman's Cultural Transmission and Evolution: A Quantitative Approach.^[116] Borrowing heavily from population genetics and epidemiology, this book built a mathematical theory concerning the spread of cultural traits. It describes the evolutionary implications of vertical transmission, passing cultural traits from parents to offspring; oblique transmission, passing cultural traits from any member of an older generation to a younger generation; and horizontal transmission, passing traits between members of the same population.

Robert Boyd and Peter Richerson's (1985) Culture and the Evolutionary Process presents models of the evolution of social learning under different environmental conditions, the population effects of social learning, various forces of selection on cultural learning rules, different forms of biased transmission and their population-level effects, and conflicts between cultural and genetic evolution.

Along with game theory, Herbert Gintis suggested that Dual inheritance theory has potential for unifying the behavioral sciences, including economics, biology, anthropology, sociology, psychology and political science because it addresses both the genetic and cultural components of human inheritance.^[117] Laland and Brown hold a similar view.^{[citation needed]}

In psychology sub-fields

Developmental psychology

According to Paul Baltes, the benefits granted by evolutionary selection decrease with age. Natural selection has not eliminated many harmful conditions and nonadaptive characteristics that appear among older adults, such as Alzheimer disease. If it were a disease that killed 20 year-olds instead of 70 year-olds this may have been a disease that natural selection could have eliminated ages ago. Thus, unaided by evolutionary pressures against nonadaptive conditions, modern humans suffer the aches, pains, and infirmities of aging and as the benefits of evolutionary selection decrease with age, the need for culture increases.^[118]

Social psychology

As humans are a highly social species, there are many adaptive problems associated with navigating the social world (e.g., maintaining allies, managing status hierarchies, interacting with outgroup members, coordinating social activities, collective decision-making). Researchers in the emerging field of evolutionary social psychology have made many discoveries pertaining to topics traditionally studied by social psychologists, including person perception, social cognition, attitudes, altruism, emotions, group dynamics, leadership, motivation, prejudice, intergroup relations, and cross-cultural differences.^[119]^[120]^[121]^[122]

When endeavouring to solve a problem humans at an early age show determination while chimpanzees have no comparable facial expression. Researchers suspect the human determined expression evolved because when a human is determinedly working on a problem other people will frequently help.^[123]

Abnormal psychology

Adaptationist hypotheses regarding the etiology of psychological disorders are often based on analogies between physiological and psychological dysfunctions,^[124] as noted in the table below. Prominent theorists and evolutionary psychiatrists include Michael T. McGuire and Randolph M. Nesse. They, and others, suggest that mental disorders are due to the interactive effects of both nature and nurture, and often have multiple contributing causes.^[12]

Possible Causes of Psychological 'Abnormalities' from an Adaptationist Perspective
Causal mechanism of failure or malfunction of adaptation	Physiological Example	Hypothesized Psychological Example
Functioning adaptation (adaptive defense)	Fever / Vomiting (functional responses to infection or ingestion of toxins)	Mild depression or anxiety (functional responses to mild loss or stress^[127]/ reduction of social interactions to prevent infection by contagious pathogens)^[128]
By-product of an adaptation(s)	Intestinal gas (byproduct of digestion of fiber)	Sexual fetishes (?) (possible byproduct of normal sexual arousal adaptations that have 'imprinted' on unusual objects or situations)
Adaptations with multiple effects	Gene for malaria resistance, in homozygous form, causes sickle cell anemia	Adaptation(s) for high levels of creativity may also predispose schizophrenia or bi-polar disorder (adaptations with both positive and negative effects, perhaps dependent on alternate developmental trajectories)
Malfunctioning adaptation	Allergies (over-reactive immunological responses)	Autism (possible malfunctioning of theory of mind module)
Frequency-dependent morphs	The two sexes / Different blood and immune system types	Personality traits and personality disorders (may represent alternative behavioral strategies dependent on the frequency of the strategy in the population)
Mismatch between ancestral & current environments	Modern diet-related Type 2 Diabetes	More frequent modern interaction with strangers (compared to family and close friends) may predispose greater incidence of depression & anxiety
Tails of normal (bell shaped) curve	Very short or tall height	Tails of the distribution of personality traits (e.g., extremely introverted or extroverted)

Evolutionary psychologists have suggested that schizophrenia and bipolar disorder may reflect a side-effect of genes with fitness benefits, such as increased creativity.^[129] (Some individuals with bipolar disorder are especially creative during their manic phases and the close relatives of schizophrenics have been found to be more likely to have creative professions.^[129]) A 1994 report by the American Psychiatry Association found that people suffered from schizophrenia at roughly the same rate in Western and non-Western cultures, and in industrialized and pastoral societies, suggesting that schizophrenia is not a disease of civilization nor an arbitrary social invention.^[129] Sociopathy may represent an evolutionarily stable strategy, by which a small number of people who cheat on social contracts benefit in a society consisting mostly of non-sociopaths.^[12] Mild depression may be an adaptive response to withdraw from, and re-evaluate, situations that have led to disadvantageous outcomes (the "analytical rumination hypothesis") ^[127] (see Evolutionary approaches to depression).

Some of these speculations have yet to be developed into fully testable hypotheses, and a great deal of research is required to confirm their validity.^[130]^[131]

Psychology of religion

Adaptationist perspectives on religious belief suggest that, like all behavior, religious behaviors are a product of the human brain. As with all other organ functions, cognition's functional structure has been argued to have a genetic foundation, and is therefore subject to the effects of natural selection and sexual selection. Like other organs and tissues, this functional structure should be universally shared amongst humans and should have solved important problems of survival and reproduction in ancestral environments. However, evolutionary psychologists remain divided on whether religious belief is more likely a consequence of evolved psychological adaptations,^[132] or is a byproduct of other cognitive adaptations.^[133]

Reception

Critics of evolutionary psychology accuse it of promoting genetic determinism, panadaptionism (the idea that all behaviors and anatomical features are adaptations), unfalsifiable hypotheses, distal or ultimate explanations of behavior when proximate explanations are superior, and malevolent political or moral ideas.^[134]

Ethical implications

Critics have argued that evolutionary psychology might be used to justify existing social hierarchies and reactionary policies.^[135]^[136] It has also been suggested by critics that evolutionary psychologists' theories and interpretations of empirical data rely heavily on ideological assumptions about race and gender.^[137]

In response to such criticism, evolutionary psychologists often caution against committing the naturalistic fallacy – the assumption that "what is natural" is necessarily a moral good.^[136]^[138]^{[page needed]}^[139] However, their caution against committing the naturalistic fallacy has been criticized as means to stifle legitimate ethical discussions.^[136]

Standard social science model

Evolutionary psychology has been entangled in the larger philosophical and social science controversies related to the debate on nature and nurture. Evolutionary psychologists typically contrast evolutionary psychology with what they call the standard social science model (SSSM). They characterize the SSSM as the "blank slate", social constructionist, or "cultural determinist" perspective that they say dominated the social sciences throughout the 20th century and assumed that the mind was shaped almost entirely by culture.^[138]
Critics have argued that evolutionary psychologists created a false dichotomy between their own view and the caricature of the SSSM.^[140]^[141]^[142] Other critics regard the SSSM as a rhetorical device or a straw man^[139]^[140]^[143] and suggest that the scientists whom evolutionary psychologists associate with the SSSM did not believe that the mind was a blank state devoid of any natural predispositions.^[139]

Reductionism and determinism

Some critics view evolutionary psychology as a form of genetic reductionism and genetic determinism,^[144]^[145] a common critique being that evolutionary psychology does not address the complexity of individual development and experience and fails to explain the influence of genes on behavior in individual cases.^[34] Evolutionary psychologists respond that EP works within a nature-nurture interactionist framework that acknowledges that many psychological adaptations are facultative (sensitive to environmental variations during individual development). EP is generally not focused on proximate analyses of behavior but rather its focus is on the study of distal/ultimate causality (the evolution of psychological adaptations). The field of behavioral genetics is focused on the study of the proximate influence of genes on behavior.^[146]

Testability of hypotheses

A frequent critique of the discipline is that the hypotheses of evolutionary psychology are frequently arbitrary and difficult or impossible to adequately test, thus questioning its status as an actual scientific discipline, for example because many current traits probably evolved to serve different functions than they do now.^[5]^[147] While evolutionary psychology hypotheses are difficult to test, evolutionary psychologists assert that it is not impossible.^[148] Part of the critique of the scientific base of evolutionary psychology includes a critique of the concept of the Environments of Evolutionary Adaptation (EEA). Some critics have argued that researchers know so little about the environment in which Homo sapiens evolved that explaining specific traits as an adaption to that environment becomes highly speculative.^[149] Evolutionary psychologists respond that they do know many things about this environment, including the facts that present day humans' ancestors were hunter-gatherers, that they generally lived in small tribes, etc.^[150]

Modularity of mind

Evolutionary psychologists generally presume that, like the body, the mind is made up of many evolved modular adaptations,^[151] although there is some disagreement within the discipline regarding the degree of general plasticity, or "generality," of some modules.^[146] It has been suggested that modularity evolves because, compared to non-modular networks, it would have conferred an advantage in terms of fitness^[152] and because connection costs are lower.^[153]
In contrast, some academics argue that it is unnecessary to posit the existence of highly domain specific modules, and, suggest that the neural anatomy of the brain supports a model based on more domain general faculties and processes.^[154]^[155] Moreover, empirical support for the domain-specific theory stems almost entirely from performance on variations of the Wason selection task which is extremely limited in scope as it only tests one subtype of deductive reasoning.^[156]^[157]

Evolutionary psychology defense

Evolutionary psychologists have addressed many of their critics (see, for example, books by Segerstråle (2000), Defenders of the Truth: The Battle for Science in the Sociobiology Debate and Beyond,^[158] Barkow (2005), Missing the Revolution: Darwinism for Social Scientists,^[159] and Alcock (2001), The Triumph of Sociobiology.^[160]).
Among their rebuttals are that some criticisms are straw men, are based on an incorrect nature versus nurture dichotomy, are based on misunderstandings of the discipline, etc.^[146]^[160]^[161]^[162]^[163]^[164]^[165]^[166]^[167] Robert Kurzban suggested that "...critics of the field, when they err, are not slightly missing the mark. Their confusion is deep and profound. It’s not like they are marksmen who can’t quite hit the center of the target; they’re holding the gun backwards."^[168]

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Thursday, May 28, 2015

Classical electromagnetism and special relativity

Transformation of the fields between inertial frames

The E and B fields

The D and H fields

The φ and A fields

The ρ and J fields

Non-relativistic approximations

Relationship between electricity and magnetism

Deriving magnetism from electrostatics

Fields intermix in different frames

Moving magnet and conductor problem

Covariant formulation in vacuum

Field tensor and 4-current

Maxwell's equations in tensor form

4-potential[edit]

How spacetime is built by quantum entanglement

Molecular evolution

Forces in molecular evolution

Mutation

Recombination

Gene conversion

Genetic drift

Selection

Genome architecture

Genome size

Repetitive elements

Chromosome number and organization

Gene content and distribution

Organelles

Origins of new genes

Molecular phylogenetics

The driving forces of evolution

Discordance with morphological evolution

Journals and societies

Evolutionary psychology

Scope

Principles

Premises

History

Theoretical foundations

Evolved psychological mechanisms

Products of evolution: adaptations, exaptations, byproducts, and random variation

Obligate and facultative adaptations

Cultural universals

Environment of evolutionary adaptedness

Mismatches

Research methods

Major areas of research

Survival and individual level psychological adaptations

Consciousness

Sensation and perception

Learning and facultative adaptations

Emotion and motivation

Cognition

Personality

Language

Mating

Parenting

Family and kin

Interactions with non-kin / reciprocity

Strong reciprocity (or "tribal reciprocity")

Evolution and culture

In psychology sub-fields

Developmental psychology

Social psychology

Abnormal psychology

Psychology of religion

Reception

Ethical implications

Standard social science model

Reductionism and determinism

Testability of hypotheses

Modularity of mind

Evolutionary psychology defense

Delayed-choice quantum eraser