Heredity

Heredity is the passing of traits to offspring (from its parent or ancestors). This is the process by which an offspring cell or organism acquires or becomes predisposed to the characteristics of its parent cell or organism. Through heredity, variations exhibited by individuals can accumulate and cause some species to evolve. The study of heredity inbiology is called genetics, which includes the field of epigenetics.

Overview

In humans, eye color is an inherited characteristic and an individual might inherit the "brown-eye trait" from one of the parents.^[1] Inherited traits are controlled by genes and the complete set of genes within an organism's genome is called its genotype.^[2]

The complete set of observable traits that make up the structure and behaviour of an organism is called its phenotype. These traits come from the interaction of its genotype with the environment.^[3] As a result, many aspects of an organism's phenotype are not inherited. For example,suntanned skin comes from the interaction between a person's genotype and sunlight; thus, suntans are not passed on to people's children. However, some people tan more easily than others, due to differences in their genotype; a striking example are people with the inherited trait ofalbinism, who do not tan at all and are very sensitive to sunburn.^[4]

Heritable traits are known to be passed from one generation to the next via DNA, a molecule that encodes genetic information.^[2] DNA is a longpolymer composed of four types of bases. The sequence of bases along a particular DNA molecule specify the genetic information, in a manner similar to a sequence of letters spelling out a sentence. Before a cell divides, the DNA is copied, so that each of the resulting two cells will inherit the DNA sequence. Portions of a DNA molecule that specify a single functional unit are called genes; different genes have different sequences of bases. Within cells, the long strands of DNA form condensed structures called chromosomes. The specific location of a DNA sequence within a chromosome is known as a locus. If the DNA sequence at a locus varies between individuals, the different forms of this sequence are called alleles. DNA sequences can change through mutations, producing new alleles. If a mutation occurs within a gene, the new allele may affect the trait that the gene controls, altering the phenotype of the organism.^[5]

However, while this simple correspondence between an allele and a trait works in some cases, most traits are more complex and are controlled by multiple interacting genes within and among organisms.^[6][7] Developmental biologists suggest that complex interactions in genetic networks and communication among cells can lead to heritable variations that may underlay some of the mechanics in developmental plasticity and canalization.^[8]

Recent findings have confirmed important examples of heritable changes that cannot be explained by direct agency of the DNA molecule. These phenomena are classed as epigenetic inheritance systems that are causally or independently evolving over genes. Research into modes and mechanisms of epigenetic inheritance is still in its scientific infancy, however, this area of research has attracted much recent activity as it broadens the scope of heritability and evolutionary biology in general.^[9] DNA methylation marking chromatin, self-sustaining metabolic loops, gene silencing by RNA interference, and the three dimensional conformation of proteins (such as prions) are areas where epigenetic inheritance systems have been discovered at the organismic level.^[10][11] Heritability may also occur at even larger scales. For example, ecological inheritance through the process of niche construction is defined by the regular and repeated activities of organisms in their environment. This generates a legacy of effect that modifies and feeds back into the selection regime of subsequent generations. Descendants inherit genes plus environmental characteristics generated by the ecological actions of ancestors.^[12] Other examples of heritability in evolution that are not under the direct control of genes include the inheritance of cultural traits, group heritability, and symbiogenesis.^[13][14][15] These examples of heritability that operate above the gene are covered broadly under the title of multilevel or hierarchical selection, which has been a subject of intense debate in the history of evolutionary science.^[14][16]

Relation to theory of evolution

When Charles Darwin proposed his theory of evolution in 1859, one of its major problems was the lack of an underlying mechanism for heredity. Darwin believed in a mix of blending inheritance and the inheritance of acquired traits (pangenesis). Blending inheritance would lead to uniformity across populations in only a few generations and thus would remove variation from a population on which natural selection could act. This led to Darwin adopting some Lamarckian ideas in later editions of On the Origin of Species and his later biological works. Darwin's primary approach to heredity was to outline how it appeared to work (noticing that traits could be inherited which were not expressed explicitly in the parent at the time of reproduction, that certain traits could be sex-linked, etc.) rather than suggesting mechanisms.

Darwin's initial model of heredity was adopted by, and then heavily modified by, his cousin Francis Galton, who laid the framework for the biometric school of heredity. Galton rejected the aspects of Darwin's pangenesis model which relied on acquired traits.

The inheritance of acquired traits was shown to have little basis in the 1880s when August Weismann cut the tails off many generations of mice and found that their offspring continued to develop tails.

History

The ancients had a variety of ideas about heredity: Theophrastus proposed that male flowers caused female flowers to ripen; Hippocrates speculated that "seeds" were produced by various body parts and transmitted to offspring at the time of conception, and Aristotle thought that male and female semen mixed at conception. Aeschylus, in 458 BC, proposed the male as the parent, with the female as a "nurse for the young life sown within her."^{[citation needed]}

Various hereditary mechanisms were envisaged without being properly tested or quantified. These included blending inheritance and the inheritance of acquired traits. Nevertheless, people were able to develop domestic breeds of animals as well as crops through artificial selection. The inheritance of acquired traits also formed a part of early Lamarckian ideas on evolution.

During the 18th century, Dutch microscopist Antonie van Leeuwenhoek (1632–1723) discovered "animalcules" in the sperm of humans and other animals. Some scientists speculated they saw a "little man" (homunculus) inside each sperm. These scientists formed a school of thought known as the "spermists." They contended the only contributions of the female to the next generation were the womb in which the homunculus grew, and prenatal influences of the womb. An opposing school of thought, the ovists, believed that the future human was in the egg, and that sperm merely stimulated the growth of the egg. Ovists thought women carried eggs containing boy and girl children, and that the gender of the offspring was determined well before conception.

Gregor Mendel: father of genetics

The idea of particulate inheritance of genes can be attributed to the Moravian^[17] monk Gregor Mendel who published his work on pea plants in 1865. However, his work was not widely known and was rediscovered in 1901. It was initially assumed the Mendelian inheritance only accounted for large (qualitative) differences, such as those seen by Mendel in his pea plants—and the idea of additive effect of (quantitative) genes was not realised until R.A. Fisher's (1918) paper, "The Correlation Between Relatives on the Supposition of Mendelian Inheritance."

Modern development of genetics and heredity

Main articles: History of genetics and Modern evolutionary synthesis

In the 1930s, work by Fisher and others resulted in a combination of Mendelian and biometric schools into the modern evolutionary synthesis. The modern synthesis bridged the gap between experimental geneticists and naturalists; and between both and palaeontologists, stating that:^[18][19]

1. All evolutionary phenomena can be explained in a way consistent with known genetic mechanisms and the observational evidence of naturalists.
2. Evolution is gradual: small genetic changes, recombination ordered by natural selection. Discontinuities amongst species (or other taxa) are explained as originating gradually through geographical separation and extinction (not saltation).
3. Selection is overwhelmingly the main mechanism of change; even slight advantages are important when continued. The object of selection is thephenotype in its surrounding environment. The role of genetic drift is equivocal; though strongly supported initially by Dobzhansky, it was downgraded later as results from ecological genetics were obtained.
4. The primacy of population thinking: the genetic diversity carried in natural populations is a key factor in evolution. The strength of natural selection in the wild was greater than expected; the effect of ecological factors such as niche occupation and the significance of barriers to gene flow are all important.
5. In palaeontology, the ability to explain historical observations by extrapolation from micro to macro-evolution is proposed. Historical contingency means explanations at different levels may exist. Gradualism does not mean constant rate of change.

The idea that speciation occurs after populations are reproductively isolated has been much debated. In plants, polyploidy must be included in any view of speciation. Formulations such as 'evolution consists primarily of changes in the frequencies of alleles between one generation and another' were proposed rather later. The traditional view is that developmental biology ('evo-devo') played little part in the synthesis, but an account of Gavin de Beer's work by Stephen Jay Gould suggests he may be an exception.^[20]

Almost all aspects of the synthesis have been challenged at times, with varying degrees of success. There is no doubt, however, that the synthesis was a great landmark in evolutionary biology. It cleared up many confusions, and was directly responsible for stimulating a great deal of research in the post-World War II era.

Trofim Lysenko however caused a backlash of what is now called Lysenkoism in the Soviet Union when he emphasised Lamarckian ideas on the inheritance of acquired traits. This movement affected agricultural research and led to food shortages in the 1960s and seriously affected the USSR.

Types of heredity

Dominant and recessive

An allele is said to be dominant if it is always expressed in the appearance of an organism (phenotype). For example, in peas the allele for green pods, G, is dominant to that for yellow pods, g. Since the allele for green pods is dominant, pea plants with the pair of alleles GG (homozygote) or Gg (heterozygote) will have green pods. The allele for yellow pods is recessive. The effects of this allele are only seen when it is present in both chromosomes, gg (homozygote).

The description of a mode of biological inheritance consists of three main categories:

1. Number of involved loci

• Monogenetic (also called "simple") – one locus
• Oligogenetic – few loci
• Polygenetic – many loci

2. Involved chromosomes

• Autosomal – loci are not situated on a sex chromosome
• Gonosomal – loci are situated on a sex chromosome
  • X-chromosomal – loci are situated on the X-chromosome (the more common case)
  • Y-chromosomal – loci are situated on the Y-chromosome
• Mitochondrial – loci are situated on the mitochondrial DNA

3. Correlation genotype–phenotype

• Dominant
• Intermediate (also called "codominant")
• Recessive

These three categories are part of every exact description of a mode of inheritance in the above order. Additionally, more specifications may be added as follows:

4. Coincidental and environmental interactions

• Penetrance
  • Complete
  • Incomplete (percentual number)
• Expressivity
  • Invariable
  • Variable
• Heritability (in polygenetic and sometimes also in oligogenetic modes of inheritance)
• Maternal or paternal imprinting phenomena (also see epigenetics)

5. Sex-linked interactions

• Sex-linked inheritance (gonosomal loci)
• Sex-limited phenotype expression (e.g., cryptorchism)
• Inheritance through the maternal line (in case of mitochondrial DNA loci)
• Inheritance through the paternal line (in case of Y-chromosomal loci)

6. Locus–locus interactions

• Epistasis with other loci (e.g., overdominance)
• Gene coupling with other loci (also see crossing over)
• Homozygotous lethal factors
• Semi-lethal factors

Determination and description of a mode of inheritance is primarily achieved through statistical analysis of pedigree data. In case the involved loci are known, methods of molecular genetics can also be employed.

Meteor shower

A meteor shower is a celestial event in which a number of meteors are observed to radiate from one point in the night sky. These meteors are caused by streams of cosmic debris called meteoroidsentering Earth's atmosphere at extremely high speeds on parallel trajectories. Most meteors are smaller than a grain of sand, so almost all of them disintegrate and never hit the Earth's surface. Intense or unusual meteor showers are known as meteor outbursts and meteor storms, which may produce greater than 1,000 meteors an hour.^[1]

The radiant point

Because meteor shower particles are all traveling in parallel paths, and at the same velocity, they will all appear to an observer below to radiate away from a single point in the sky. This radiant point is caused by the effect of perspective, similar to railroad tracks converging at a single vanishing point on the horizon when viewed from the middle of the tracks. Meteor showers are almost always named after the constellation from which the meteors appear to originate. This "fixed point" slowly moves across the sky during the night due to the Earth turning on its axis, the same reason the stars appear to slowly march across the sky. The radiant also moves slightly from night to night against the background stars (radiant drift) due to the Earth moving in its orbit around the sun. See "IMO" Meteor Shower Calendar 2007 (International Meteor Organization) for maps of drifting "fixed points."

Meteor showers are named after the nearest bright star with a Greek or Roman letter assigned that is close to the radiant position at the peak of the shower, whereby the declension of the Latin possessive form is replaced by "id" or "ids". Hence, meteors radiating from near the star delta Aquarii (declension "-i") are called delta Aquariids. The International Astronomical Union's Task Group on Meteor Shower Nomenclature and the IAU's Meteor Data Center keep track of meteor shower nomenclature and which showers are established.

The origin of meteoroid streams

Comet Encke's meteoroid trail is the diagonal red glow

Meteoroid trail between fragments ofComet 73P

A meteor shower is the result of an interaction between a planet, such as Earth, and streams of debris from a comet. Comets can produce debris by water vapor drag, as demonstrated by Fred Whipple in 1951,^[2] and by breakup. Whipple envisioned comets as "dirty snowballs," made up of rock embedded in ice, orbiting the Sun. The "ice" may be water, methane, ammonia, or other volatiles, alone or in combination. The "rock" may vary in size from that of a dust mote to that of a small boulder. Dust mote sized solids are orders of magnitude more common than those the size of sand grains, which, in turn, are similarly more common than those the size of pebbles, and so on. When the ice warms and sublimates, the vapor can drag along dust, sand, and pebbles. Each time a comet swings by the Sun in its orbit, some of its ice vaporizes and a certain amount of meteoroids will be shed. The meteoroids spread out along the entire orbit of the comet to form a meteoroid stream, also known as a "dust trail" (as opposed to a comet's "dust tail" caused by the very small particles that are quickly blown away by solar radiation pressure).

Recently, Peter Jenniskens ^[3] has argued that most of our short-period meteor showers are not from the normal water vapor drag of active comets, but the product of infrequent disintegrations, when large chunks break off a mostly dormant comet. Examples are the Quadrantids and Geminids, which originated from a breakup of asteroid-looking objects 2003 EH1 and 3200 Phaethon, respectively, about 500 and 1000 years ago. The fragments tend to fall apart quickly into dust, sand, and pebbles, and spread out along the orbit of the comet to form a dense meteoroid stream, which subsequently evolves into Earth's path.

The dynamical evolution of meteoroid streams

Shortly after Whipple predicted that dust particles travelled at low speeds relative to the comet, Milos Plavec was the first to offer the idea of a dust trail, when he calculated how meteroids, once freed from the comet, would drift mostly in front of or behind the comet after completing one orbit. The effect is simpleorbital mechanics - the material drifts only a little laterally away from the comet while drifting ahead or behind the comet because some particles make a wider orbit than others.^[3] These dust trails are sometimes observed in comet images taken at mid infrared wavelengths (heat radiation), where dust particles from the previous return to the Sun are spread along the orbit of the comet (see figures).

The gravitational pull of the planets determines where the dust trail would pass by Earth orbit, much like a gardener directing a hose to water a distant plant. Most years, those trails would miss the Earth altogether, but in some years the Earth is showered by meteors. This effect was first demonstrated from observations of the 1995 alpha Monocerotids,^[4][5] and from earlier not widely known identifications of past earth storms.

In the 1890s, Irish astronomer George Johnstone Stoney (1826–1911) and British astronomer Arthur Matthew Weld Downing (1850–1917), were the first to attempt to calculate the position of the dust at Earth's orbit. They studied the dust ejected in 1866 by comet 55P/Tempel-Tuttle in advance of the anticipated Leonid shower return of 1898 and 1899. Meteor storms were anticipated, but the final calculations showed that most of the dust would be far inside of Earth's orbit. The same results were independently arrived at by Adolf Berberich of the Königliches Astronomisches Rechen Institut (Royal Astronomical Computation Institute) in Berlin, Germany. Although the absence of meteor storms that season confirmed the calculations, the advance of much better computing tools was needed to arrive at reliable predictions.

In 1985, E. D. Kondrat'eva and E. A. Reznikov of Kazan State University first correctly identified the years when dust was released responsible for several past Leonid meteor storms. In anticipation of the 1999 Leonid storm, Robert H. McNaught^[6] David Asher,^[7] and Finland's Esko Lyytinen were the first to apply this method in the West.^[8][9] Peter Jenniskens has published predictions for future dust trail encounters, resulting in "meteor storms" or "meteor outbursts" for the next 50 years.^[3]

Over longer periods of time, the dust trails can evolve in complicated ways. One effect is that the orbits of some repeating comets, and meteoroids leaving them, are in resonant orbits with Jupiter or one of the other large planets - so many revolutions of one will equal another number of revolutions of the other. So over time since Jupiter will have the same relative position intermittently and it will tend to pull meteoroids into keeping that relative position. This creates a shower component called a "filament."

A second effect is a close encounter with a planet. When the meteoroids pass by Earth, some are accelerated (making wider orbits around the Sun), others are decelerated (making shorter orbits), resulting in gaps in the dust trail in the next return (like opening a curtain, with grains piling up at the beginning and end of the gap). Also, Jupiter's perturbation can change sections of the dust trail dramatically, especially for short period comets, when the grains approach the big planet at their furthest point along the orbit around the Sun, moving most slowly. As a result, the trail has a clumping, a braiding or a tangling of crescents, of each individual release of material.

The third effect is that of radiation pressure which will push less massive particles into orbits further from the sun - while more massive objects (responsible for bolides or fireballs) will tend to be affected less by radiation pressure. This makes some dust trail encounters rich in bright meteors, others rich in faint meteors. Over time, these effects disperse the meteoroids and create a broader stream. The meteors we see from these streams are part of annual showers, because Earth encounters those streams every year at much the same rate.

When the meteoroids collide with other meteoroids in the zodiacal cloud, they lose their stream association and become part of the "sporadic meteors" background. Long since dispersed from any stream or trail, they form isolated meteors, not a part of any shower. These random meteors will not appear to come from the radiant of the main shower.

Notable meteor showers

Perseid and Leonid meteor showers

The most visible meteor shower in most years are the Perseids, which peak on August 12 of each year at over 1 meteor a minute. A useful tool to calculate how many meteors per hour are visible from your observing location is found here: http://leonid.arc.nasa.gov/estimator.html.

The most spectacular meteor shower is probably the Leonids, the King of Meteor Showers^[10] which peaks on a day near 17 November. Approximately every 33 years the Leonid shower produces a "meteor storm", peaking at rates of thousands of meteors per hour. These Leonid storms gave birth to the term "meteor shower", when it was first realised during the November 1833 storm that the meteors radiated from near the star Gamma Leonis. The last Leonid storms were in 1999, 2001 (two), and 2002 (two). Before that, there were storms in 1767, 1799, 1833, 1866, 1867, and 1966. When the Leonid shower is not storming it is less active than the Perseids.

Other notable meteor showers

Further information: List of meteor showers

Official names are given in the International Astronomical Union meteor shower list.^[11]

Shower	Time	Parent object
Quadrantids	Early January	The same as the parent object of minor planet 2003 EH1,[12] and perhaps comets C/1490 Y1 and C/1385 U1 [13]
Lyrids	late April	Comet Thatcher
Pi Puppids (periodic)	late April	Comet 26P/Grigg-Skjellerup
Eta Aquariids	early May	Comet 1P/Halley
Arietids	mid June	Comet 96P/Machholz, Marsden and Kracht comet groups complex [1][14]
June Bootids (periodic)	late June	Comet 7P/Pons-Winnecke
Southern Delta Aquariids	late July	Comet 96P/Machholz, Marsden and Kracht comet groups complex [1][14]
Alpha Capricornids	late July	Comet 169P/NEAT[15]
Perseids	mid-August	Comet 109P/Swift-Tuttle
Kappa Cygnids	mid-August	Minor planet 2008 ED69[16]
Aurigids (periodic)	early September	Comet C/1911 N1 (Kiess)[17]
Giacobinids (periodic)	early October	Comet 21P/Giacobini-Zinner
Orionids	late October	Comet 1P/Halley
Southern Taurids	early November	Comet 2P/Encke
Northern Taurids	mid-November	Minor planet 2004 TG10 and others [1][18]
Andromedids (periodic)	mid-November	Comet 3D/Biela[19]
Alpha Monocerotids (periodic)	mid-November	unknown [20]
Leonids	mid-November	Comet 55P/Tempel-Tuttle
Phoenicids (periodic)	early-December	Comet D/1819 W1 (Blanpain)[21]
Geminids	mid-December	Minor planet 3200 Phaethon[22]
Ursids	late December	Comet 8P/Tuttle[23]

Extraterrestrial meteor showers

Any other solar system body with a reasonably transparent atmosphere can also have meteor showers. For instance, Mars is known to have meteor showers,^[24] although these are different from the ones seen on Earth because the different orbits of Mars and Earth intersect orbits of comets in different ways.

Although the Martian atmosphere has less than one percent of the density of Earth's at ground level, at their upper edges, where meteoroids strike, the two are more similar. Because of the similar air pressure at altitudes for meteors, the effects are much the same. Only the relatively slower motion of the meteoroids due to increased distance from the sun should marginally decrease meteor brightness. This is somewhat balanced in that the slower descent means that Martian meteors have more time in which to ablate.^[25]

On March 7, 2004, the panoramic camera on Mars Exploration Rover Spirit recorded a streak which is now believed to have been caused by a meteor from a Martian meteor shower associated with comet 114P/Wiseman-Skiff. A strong display from this shower was expected on December 20, 2007. Other showers speculated about are a "Lambda Geminid" shower associated with the Eta Aquariids of Earth (i.e. both associated with Comet 1P/Halley), a "Beta Canis Major" shower associated with Comet 13P/Olbers, and "Draconids" from 5335 Damocles.^[26]