ratio of word probabilities predicted from brain for eye and butterfly

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eye

butterfly

top 10 words in brain distribution (in article):
water form light animal time surface type cause region land
top 10 words in brain distribution (in article):
species breed male human female size kill range live common
top 10 words in brain distribution (not in article):
ice rock drink river flow sea lamp ocean soil wind
top 10 words in brain distribution (not in article):
animal cat wolf hunt dog wild population bear material lion
times more probable under eye 30 20 10 6 4 2.5 1.25 1 1.25 2.5 4 6 10 20 30 times more probable under butterfly
(words not in the model)
Eyes'" are organs that detect light, and send signals along the optic nerve to the visual and other areas of the brain. Complex optical systems with resolving power have come in ten fundamentally different forms, and 96% of animal species possess a complex optical system. Image-resolving eyes are present in cnidaria, mollusks, chordates, annelids and arthropods. The simplest "eyes", in even unicellular organisms, do nothing but detect whether the surroundings are light or dark, which is sufficient for the entrainment of circadian rhythms. From more complex eyes, retinal photosensitive ganglion cells send signals along the retinohypothalamic tract to the suprachiasmatic nuclei to effect circadian adjustment. Overview. Complex eyes can distinguish shapes and colors. The visual fields of many organisms, especially predators, involve large areas of binocular vision to improve depth perception; in other organisms, eyes are located so as to maximise the field of view, such as in rabbits and horses. The first proto-eyes evolved among animals 540 million years ago, about the time of the so-called Cambrian explosion. The last common ancestor of animals possessed the biochemical toolkit necessary for vision, and more advanced eyes have evolved in 96% of animal species in 6 of the thirty-something main phyla. In most vertebrates and some mollusks, the eye works by allowing light to enter it and project onto a light-sensitive panel of cells, known as the retina, at the rear of the eye. The cone cells (for color) and the rod cells (for low-light contrasts) in the retina detect and convert light into neural signals for vision. The visual signals are then transmitted to the brain via the optic nerve. Such eyes are typically roughly spherical, filled with a transparent gel-like substance called the vitreous humour, with a focusing lens and often an iris; the relaxing or tightening of the muscles around the iris change the size of the pupil, thereby regulating the amount of light that enters the eye, and reducing aberrations when there is enough light. The eyes of cephalopods, fish, amphibians and snakes usually have fixed lens shapes, and focusing vision is achieved by telescoping the lens similar to how a camera focuses. Compound eyes are found among the arthropods and are composed of many simple facets which, depending on the details of anatomy, may give either a single pixelated image or multiple images, per eye. Each sensor has its own lens and photosensitive cell(s). Some eyes have up to 28,000 such sensors, which are arranged hexagonally, and which can give a full 360-degree field of vision. Compound eyes are very sensitive to motion. Some arthropods, including many Strepsiptera, have compound eyes of only a few facets, each with a retina capable of creating an image, creating vision. With each eye viewing a different thing, a fused image from all the eyes is produced in the brain, providing very different, high-resolution images. Possessing detailed hyperspectral color vision, the Mantis shrimp has been reported to have the world's most complex color vision system. Trilobites, which are now extinct, had unique compound eyes. They used clear calcite crystals to form the lenses of their eyes. In this, they differ from most other arthropods, which have soft eyes. The number of lenses in such an eye varied, however: some trilobites had only one, and some had thousands of lenses in one eye. In contrast to compound eyes, simple eyes are those that have a single lens. For example, jumping spiders have a large pair of simple eyes with a narrow field of view, supported by an array of other, smaller eyes for peripheral vision. Some insect larvae, like caterpillars, have a different type of simple eye (stemmata) which gives a rough image. Some of the simplest eyes, called ocelli, can be found in animals like some of the snails, which cannot actually "see" in the normal sense. They do have photosensitive cells, but no lens and no other means of projecting an image onto these cells. They can distinguish between light and dark, but no more. This enables snails to keep out of direct sunlight. In organisms dwelling near deep-sea vents, compound eyes have been secondarily simplified and adapted to spot the infra-red light produced by the hot vents in this way the bearers can spot hot springs and avoid being boiled alive. Evolution. Visual pigments appear to have a common ancestor and were probably involved in circadian rhythms or reproductive timing in simple organisms. Complex vision, associated with dedicated visual organs, or eyes, evolved many times in different lineages. Types of eye. Nature has produced ten different eye layouts indeed every way of capturing an image has evolved at least once in nature, with the exception of zoom and Fresnel lenses. Eye types can be categorized into "simple eyes", with one concave chamber, and "compound eyes", which comprise a number of individual lenses laid out on a convex surface. Note that "simple" does not imply a reduced level of complexity or acuity. Indeed, any eye type can be adapted for almost any behaviour or environment. The only limitations specific to eye types are that of resolution the physics of compound eyes prevents them from achieving a resolution better than 1°. Also, superposition eyes can achieve greater sensitivity than apposition eyes, so are better suited to dark-dwelling creatures. Eyes also fall into two groups on the basis of their photoreceptor's cellular construction, with the photoreceptor cells either being cilliated (as in the vertebrates) or rhabdomic. These two groups are not monophyletic; the cnidaira also possess cilliated cells, Pit eyes. Pit eyes, also known as stemma, are eye-spots which may be set into a pit to reduce the angles of light that enters and affects the eyespot, to allow the organism to deduce the angle of incoming light. Found in about 85% of phyla, these basic forms were probably the precursors to more advanced types of "simple eye". They are small, comprising up to about 100 cells covering about 100 µm. The directionality can be improved by reducing the size of the aperture, by incorporating a reflective layer behind the receptor cells, or by filling the pit with a refractile material. Pinhole eye. The pinhole eye is an "advanced" form of pit eye incorporating these improvements, most notably a small aperture (which may be adjustable) and deep pit. It is only found in the nautiloids. Without a lens to focus the image, it produces a blurry image, and will blur out a point to the size of the aperture. Consequently, nautiloids can't discriminate between objects with an angular separation of less than 11°. Shrinking the aperture would produce a sharper image, but let in less light. Spherical lensed eye. The resolution of pit eyes can be greatly improved by incorporating a material with a higher refractive index to form a lens, which may greatly reduce the blur radius encountered hence increasing the resolution obtainable. The most basic form, still seen in some gastropods and annelids, consists of a lens of one refractive index. A far sharper image can be obtained using materials with a high refractive index, decreasing to the edges this decreases the focal length and thus allows a sharp image to form on the retina. This also allows a larger aperture for a given sharpness of image, allowing more light to enter the lens; and a flatter lens, reducing spherical aberration. Such an inhomogeneous lens is necessary in order for the focal length to drop from about 4 times the lens radius, to 2.5 radii. Heterogeneous eyes have evolved at least eight times four or more times in gastropods, once in the copepods, once in the annelids and once in the cephalopods. No aquatic organisms possess homogeneous lenses; presumably the evolutionary pressure for a heterogeneous lens is great enough for this stage to be quickly "outgrown". This eye creates an image that is sharp enough that motion of the eye can cause significant blurring. To minimize the effect of eye motion while the animal moves, most such eyes have stabilizing eye muscles. The ocelli of insects bear a simple lens, but their focal point always lies behind the retina; consequently they can never form a sharp image. This capitulates the function of the eye. Ocelli (pit-type eyes of arthropods) blur the image across the whole retina, and are consequently excellent at responding to rapid changes in light intensity across the whole visual field this fast response is further accelerated by the large nerve bundles which rush the information to the brain. Focussing the image would also cause the sun's image to be focussed on a few receptors, with the possibility of damage under the intense light; shielding the receptors would block out some light and thus reduce their sensitivity. This fast response has led to suggestions that the ocelli of insects are used mainly in flight, because they can be used to detect sudden changes in which way is up (because light, especially UV light which is absorbed by vegetation, usually comes from above). Weaknesses. One weakness of this eye construction is that chromatic aberration is still quite high although for organisms without color vision, this is a very minor concern. A weakness of the vertebrate eye is the blind spot which results from a gap in the retina where the optic nerve exits at the back of the eye; the cephalopod eye has no blind spot as the retina is in the opposite orientation. Multiple lenses. Some marine organisms bear more than one lens; for instance the copeopod "Pontella" has three. The outer has a parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed. "Copilla'"s eyes have two lenses, which move in and out like a telescope. Such arrangements are rare and poorly understood, but represent an interesting alternative construction. An interesting use of multiple lenses is seen in some hunters such as eagles and jumping spiders, which have a refractive cornea (discussed next): these have a negative lens, enlarging the observed image by up to 50% over the receptor cells, thus increasing their optical resolution. Refractive cornea. In the eyes of most terrestrial vertebrates (along with spiders and some insect larvae) the vitreous fluid has a higher refractive index than the air, relieving the lens of the function of reducing the focal length. This has freed it up for fine adjustments of focus, allowing a very high resolution to be obtained. As with spherical lenses, the problem of spherical aberration caused by the lens can be countered either by using an inhomogeneous lens material, or by flattening the lens. Flattening the lens has a disadvantage: the quality of vision is diminished away from the main line of focus, meaning that animals requiring all-round vision are detrimented. Such animals often display an inhomogeneous lens instead. As mentioned above, a refractive cornea is only useful out of water; in water, there is no difference in refractive index between the vitreous fluid and the surrounding water. Hence creatures which have returned to the water penguins and seals, for example lose their refractive cornea and return to lens-based vision. An alternative solution, borne by some divers, is to have a very strong cornea. Reflector eyes. An alternative to a lens is to line the inside of the eye with mirrors", and reflect the image to focus at a central point. The nature of these eyes means that if one were to peer into the pupil of an eye, one would see the same image that the organism would see, reflected back out. Many small organisms such as rotifers, copeopods and platyhelminths use such organs, but these are too small to produce usable images. Some larger organisms, such as scallops, also use reflector eyes. The scallop "Pecten" has up to 100 millimeter-scale reflector eyes fringing the edge of its shell. It detects moving objects as they pass successive lenses. Compound eyes. A compound eye may consist of thousands of individual photoreception units. The image perceived is a combination of inputs from the numerous ommatidia (individual "eye units"), which are located on a convex surface, thus pointing in slightly different directions. Compared with simple eyes, compound eyes possess a very large view angle, and can detect fast movement and, in some cases, the polarization of light. Because the individual lenses are so small, the effects of diffraction impose a limit on the possible resolution that can be obtained. This can only be countered by increasing lens size and number to see with a resolution comparable to our simple eyes, humans would require compound eyes which would each reach the size of their head. Compound eyes fall into two groups: apposition eyes, which form multiple inverted images, and superposition eyes, which form a single erect image. Compound eyes are common in arthropods, and are also present in annelids and some bivalved molluscs. Compound eyes, in arthropods at least, grow at their margins by the addition of new ommatidia. Apposition eyes. Apposition eyes are the most common form of eye, and are presumably the ancestral form of compound eye. They are found in all arthropod groups, although they may have evolved more than once within this phylum. Some annelids and bivalves also have apposition eyes. They are also possessed by "Limulus", the horseshoe crab, and there are suggestions that other chelicerates developed their simple eyes by reduction from a compound starting point. (Some caterpillars appear to have evolved compound eyes from simple eyes in the opposite fashion.) Apposition eyes work by gathering a number of images, one from each eye, and combining them in the brain, with each eye typically contributing a single point of information. The typical apposition eye has a lens focusing light from one direction on the rhabdom, while light from other directions is absorbed by the dark wall of the ommatidium. In the other kind of apposition eye, found in the Strepsiptera, lenses are not fused to one another, and each forms an entire image; these images are combined in the brain. This is called the schizochroal compound eye or the neural superposition eye. Because images are combined additively, this arrangement allows vision under lower light levels. Superposition eyes. The second type is named the superposition eye. The superposition eye is divided into three types; the refracting, the reflecting and the parabolic superposition eye. The refracting superposition eye has a gap between the lens and the rhabdom, and no side wall. Each lens takes light at an angle to its axis and reflects it to the same angle on the other side. The result is an image at half the radius of the eye, which is where the tips of the rhabdoms are. This kind is used mostly by nocturnal insects. In the parabolic superposition compound eye type, seen in arthropods such as mayflies, the parabolic surfaces of the inside of each facet focus light from a reflector to a sensor array. Long-bodied decapod crustaceans such as shrimp, prawns, crayfish and lobsters are alone in having reflecting superposition eyes, which also has a transparent gap but uses corner mirrors instead of lenses. Parabolic superposition. This eye type functions by refracting light, then using a parabolic mirror to focus the image; it combines features of superposition and apposition eyes. Other. Good fliers like flies or honey bees, or prey-catching insects like praying mantis or dragonflies, have specialized zones of ommatidia organized into a fovea area which gives acute vision. In the acute zone the eye are flattened and the facets larger. The flattening allows more ommatidia to receive light from a spot and therefore higher resolution. There are some exceptions from the types mentioned above. Some insects have a so-called single lens compound eye, a transitional type which is something between a superposition type of the multi-lens compound eye and the single lens eye found in animals with simple eyes. Then there is the mysid shrimp "Dioptromysis paucispinosa". The shrimp has an eye of the refracting superposition type, in the rear behind this in each eye there is a single large facet that is three times in diameter the others in the eye and behind this is an enlarged crystalline cone. This projects an upright image on a specialized retina. The resulting eye is a mixture of a simple eye within a compound eye. Another version is the pseudofaceted eye, as seen in Scutigera. This type of eye consists of a cluster of numerous ocelli on each side of the head, organized in a way that resembles a true compound eye. The body of "Ophiocoma wendtii", a type of brittle star, is covered with ommatidia, turning its whole skin into a compound eye. The same is true of many chitons. Relationship to lifestyle. Eyes are generally adapted to the environment and lifestyle of the organism which bears them. For instance, the distribution of photoreceptors tends to match the area in which the highest acuity is required, with horizon-scanning organisms, such as those that live on the African plains, having a horizontal line of high-density ganglia, while tree-dwelling creatures which require good all-round vision tend to have a symmetrical distribution of ganglia, with acuity decreasing outwards from the centre. Of course, for most eye types, it is impossible to diverge from a spherical form, so only the density of optical receptors can be altered. In organisms with compound eyes, it is the number of ommatidia rather than ganglia that reflects the region of highest data acquisition. Optical superposition eyes are constrained to a spherical shape, but other forms of compound eyes may deform to a shape where more ommatidia are aligned to, say, the horizon, without altering the size or density of individual ommatidia. Eyes of horizon-scanning organisms have stalks so they can be easily aligned to the horizon when this is inclined, for example if the animal is on a slope. An extension of this concept is that the eyes of predators typically have a zone of very acute vision at their centre, to assist in the identification of prey. In deep water organisms, it may not be the centre of the eye that is enlarged. The hyperiid amphipods are deep water animals that feed on organisms above them. Their eyes are almost divided into two, with the upper region thought to be involved in detecting the silhouettes of potential prey or predators against the faint light of the sky above. Accordingly, deeper water hyperiids, where the light against which the silhouettes must be compared is dimmer, have larger "upper-eyes", and may lose the lower portion of their eyes altogether. Depth perception can be enhanced by having eyes which are enlarged in one direction; distorting the eye slightly allows the distance to the object to be estimated with a high degree of accuracy. Acuity is higher among male organisms that mate in mid-air, as they need to be able to spot and assess potential mates against a very large backdrop. On the other hand, the eyes of organisms which operate in low light levels, such as around dawn and dusk or in deep water, tend to be larger to increase the amount of light that can be captured. It is not only the shape of the eye that may be affected by lifestyle. Eyes can be the most visible parts of organisms, and this can act as a pressure on organisms to have more transparent eyes at the cost of function. Eyes may be mounted on stalks to provide better all-round vision, by lifting them above an organism's carapace; this also allows them to track predators or prey without moving the head. Acuity. Visual acuity is often measured in cycles per degree (CPD), which measures an angular resolution, or how much an eye can differentiate one object from another in terms of visual angles. Resolution in CPD can be measured by bar charts of different numbers of white black stripe cycles. For example, if each pattern is 1.75 cm wide and is placed at 1 m distance from the eye, it will subtend an angle of 1 degree, so the number of white black bar pairs on the pattern will be a measure of the cycles per degree of that pattern. The highest such number that the eye can resolve as stripes, or distinguish from a gray block, is then the measurement of visual acuity of the eye. For a human eye with excellent acuity, the maximum theoretical resolution would be 50 CPD (1.2 arcminute per line pair, or a 0.35 mm line pair, at 1 m). A rat can resolve only about 1 to 2 CPD. A horse has higher acuity through most of the visual field of its eyes than a human has, but does not match the high acuity of the human eye's central fovea region. Spherical aberration limits the resolution of a 7 mm pupil to about 3 arcminutes per line pair. At a pupil diameter of 3 mm, the spherical aberration is greatly reduced, resulting in an improved resolution of approximately 1.7 arcminutes per line pair. A resolution of 2 arcminutes per line pair, equivalent to a 1 arcminute gap in an optotype, corresponds to 20 20 (normal vision) in humans. Color. All organisms are restricted to a small range of the electromagnetic spectrum; this varies from creature to creature, but is mainly between 400 and 700 nm. A butterfly'" is an insect of the order Lepidoptera. Like all Lepidoptera, butterflies are notable for their unusual life cycle with a larval caterpillar stage, an inactive pupal stage, and a spectacular metamorphosis into a familiar and colourful winged adult form. Most species are day-flying so they regularly attract attention. The diverse patterns formed by their brightly coloured wings and their erratic yet graceful flight have made butterfly watching a hobby. Butterflies comprise the "true butterflies" (superfamily Papilionoidea), the "skippers" (superfamily Hesperioidea) and the "moth-butterflies" (superfamily Hedyloidea). Butterflies exhibit polymorphism, mimicry and aposematism. Some migrate over long distances. Some butterflies have evolved symbiotic and parasitic relationships with social insects such as ants. Butterflies are important economically as agents of pollination. In addition, a few species are pests, because they can damage domestic crops and trees in their larval stage. Culturally, butterflies are a popular motif in the visual and literary arts. The four-stage lifecycle. Unlike many insects, butterflies do not experience a nymph period, but instead go through a pupal stage which lies between the larva and the adult stage (the "imago"). Butterflies are termed as holometabolous insects, and go through complete metamorphosis. It is a popular belief that butterflies have very short life spans. However, butterflies in their adult stage can live from a week to nearly a year depending on the species. Many species have long larval life stages while others can remain dormant in their pupal or egg stages and thereby survive winters. Butterflies may have one or more broods per year. The number of generations per year varies from temperate to tropical regions with tropical regions showing a trend towards multivoltinism. Egg. Butterfly eggs consist of a hard-ridged outer layer of shell, called the "chorion". This is lined with a thin coating of wax which prevents the egg from drying out before the larva has had time to fully develop. Each egg contains a number of tiny funnel-shaped openings at one end, called "micropyles"; the purpose of these holes is to allow sperm to enter and fertilize the egg. Butterfly and moth eggs vary greatly in size between species, but they are all either spherical or ovate. Butterfly eggs are fixed to a leaf with a special glue which hardens rapidly. As it hardens it contracts, deforming the shape of the egg. This glue is easily seen surrounding the base of every egg forming a meniscus. The nature of the glue is unknown and is a suitable subject for research. The same glue is produced by a pupa to secure the setae of the cremaster. This glue is so hard that the silk pad, to which the setae are glued, cannot be separated. Eggs are usually laid on plants. Each species of butterfly has its own hostplant range and while some species of butterfly are restricted to just one species of plant, others use a range of plant species, often including members of a common family. The egg stage lasts a few weeks in most butterflies but eggs laid close to winter, especially in temperate regions, go through a "diapause" stage, and the hatching may take place only in spring. Other butterflies may lay their eggs in the spring and have them hatch in the summer. These butterflies are usually northern species (Mourning Cloak, Tortoiseshells) Caterpillars. Butterfly larvae, or caterpillars, consume plant leaves and spend practically all of their time in search of food. Although most caterpillars are herbivorous, a few species such as "Spalgis epius" and "Liphyra brassolis" are entomophagous (insect eating). Some larvae, especially those of the Lycaenidae, form mutual associations with ants. They communicate with the ants using vibrations that are transmitted through the substrate as well as using chemical signals. The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions. Caterpillars mature through a series of stages called instars. Near the end of each instar, the larva undergoes a process called apolysis, in which the cuticle, a mixture of chitin and specialized proteins, is released from the epidermis and the epidermis begins to form a new cuticle beneath. At the end of each instar, the larva moults the old cuticle, and the new cuticle rapidly hardens and pigments. Development of butterfly wing patterns begins by the last larval instar. Butterfly caterpillars have three pairs of true legs from the thoracic segments and up to 6 pairs of prolegs arising from the abdominal segments. These prolegs have rings of tiny hooks called crochets that help them grip the substrate. Some caterpillars have the ability to inflate parts of their head to appear snake-like. Many have false eye-spots to enhance this effect. Some caterpillars have special structures called osmeteria which are everted to produce smelly chemicals. These are used in defense. Host plants often have toxic substances in them and caterpillars are able to sequester these substances and retain them into the adult stage. This helps making them unpalatable to birds and other predators. Such unpalatibility is advertised using bright red, orange, black or white warning colours. The toxic chemicals in plants are often evolved specifically to prevent them from being eaten by insects. Insects in turn develop countermeasures or make use of these toxins for their own survival. This "arms race" has led to the coevolution of insects and their host plants. Wing development. Wings or wing pads are not visible on the outside of the larva, but when larvae are dissected, tiny developing "wing disks" can be found on the second and third thoracic segments, in place of the spiracles that are apparent on abdominal segments. Wing disks develop in association with a trachea that runs along the base of the wing, and are surrounded by a thin "peripodial membrane", which is linked to the outer epidermis of the larva by a tiny duct. Wing disks are very small until the last larval instar, when they increase dramatically in size, are invaded by branching tracheae from the wing base that precede the formation of the wing veins, and begin to develop patterns associated with several landmarks of the wing. Near pupation, the wings are forced outside the epidermis under pressure from the hemolymph, and although they are initially quite flexible and fragile, by the time the pupa breaks free of the larval cuticle they have adhered tightly to the outer cuticle of the pupa (in obtect pupae). Within hours, the wings form a cuticle so hard and well-joined to the body that pupae can be picked up and handled without damage to the wings. Pupa. When the larva is fully grown, hormones such as prothoracicotropic hormone (PTTH) are produced. At this point the larva stops feeding and begins "wandering" in the quest of a suitable pupation site, often the underside of a leaf. The larva transforms into a pupa (or chrysalis) by anchoring itself to a substrate and moulting for the last time. The chrysalis is usually incapable of movement, although some species can rapidly move the abdominal segments or produce sounds to scare potential predators. The pupal transformation into a butterfly through metamorphosis has held great appeal to mankind. To transform from the miniature wings visible on the outside of the pupa into large structures usable for flight, the pupal wings undergo rapid mitosis and absorb a great deal of nutrients. If one wing is surgically removed early on, the other three will grow to a larger size. In the pupa, the wing forms a structure that becomes compressed from top to bottom and pleated from proximal to distal ends as it grows, so that it can rapidly be unfolded to its full adult size. Several boundaries seen in the adult color pattern are marked by changes in the expression of particular transcription factors in the early pupa. Adult or imago. The adult, sexually mature, stage of the insect is known as the imago. As Lepidoptera, butterflies have four wings that are covered with tiny scales (see photo). The fore and hindwings are not hooked together, permitting a more graceful flight. An adult butterfly has six legs, but in the nymphalids, the first pair is reduced. After it emerges from its pupal stage, a butterfly cannot fly until the wings are unfolded. A newly-emerged butterfly needs to spend some time inflating its wings with blood and letting them dry, during which time it is extremely vulnerable to predators. Some butterflies' wings may take up to three hours to dry while others take about one hour. Most butterflies and moths will excrete excess dye after hatching. This fluid may be white, red, orange, or in rare cases, blue. External morphology. Butterflies have two antennae, two compound eyes, and a proboscis. Adult butterflies have four wings: a forewing and hindwing on both the left and the right side of the body. The body is divided into three segments: the head, thorax, and the abdomen. They have two antennae, two compound eyes, and a proboscis. Scales. Butterflies are characterized by their scale-covered wings. The coloration of butterfly wings is created by minute scales. These scales are pigmented with melanins that give them blacks and browns, but blues, greens, reds and iridescence are usually created not by pigments but the microstructure of the scales. This structural coloration is the result of coherent scattering of light by the photonic crystal nature of the scales. The scales cling somewhat loosely to the wing and come off easily without harming the butterfly. Polymorphism. Many adult butterflies exhibit polymorphism, showing differences in appearance. These variations include geographic variants and seasonal forms. In addition many species have females in multiple forms, often with mimetic forms. Sexual dimorphism in coloration and appearance is widespread in butterflies. In addition many species show sexual dimorphism in the patterns of ultraviolet reflectivity, while otherwise appearing identical to the unaided human eye. Most of the butterflies have a sex-determination system that is represented as ZW with females being the heterogametic sex (ZW) and males homogametic (ZZ). Genetic abnormalities such as gynandromorphy also occur from time to time. In addition many butterflies are infected by "Wolbachia" and infection by the bacteria can lead to the conversion of males into females or the selective killing of males in the egg stage. Mimicry. Batesian and Mullerian mimicry in butterflies is common. Batesian mimics imitate other species to enjoy the protection of an attribute they do not share, aposematism in this case. The Common Mormon of India has female morphs which imitate the unpalatable red-bodied swallowtails, the Common Rose and the Crimson Rose. Mullerian mimicry occurs when aposematic species evolve to resemble each other, presumably to reduce predator sampling rates, the Heliconius butterflies from the Americas being a good example. Wing markings called eyespots are present in some species; these may have an automimicry role for some species. In others, the function may be intraspecies communication, such as mate attraction. In several cases, however, the function of butterfly eyespots is not clear, and may be an evolutionary anomaly related to the relative elasticity of the genes that encode the spots. Seasonal polyphenism. div name="wet-dry forms" Many of the tropical butterflies have distinctive seasonal forms. This phenomenon is termed "seasonal polyphenism" and the seasonal forms of the butterflies are called the dry-season and wet-season forms. How the season affects the genetic expression of patterns is still a subject of research. Experimental modification by ecdysone hormone treatment has demonstrated that it is possible to control the continuum of expression of variation between the wet and dry-season forms. The dry-season forms are usually more cryptic and it has been suggested that the protection offered may be an adaptation. Some also show greater dark colours in the wet-season form which may have thermoregulatory advantages by increasing ability to absorb solar radiation. Habits. Butterflies feed primarily on nectar from flowers. Some also derive nourishment from pollen, tree sap, rotting fruit, dung, and dissolved minerals in wet sand or dirt. Butterflies are important as pollinators for some species of plants although in general they do not carry as much pollen load as the Hymenoptera. They are however capable of moving pollen over greater distances. Within the Lepidoptera, the Hawkmoths and the Noctuidae are dominant as pollinators. As adults, butterflies consume only liquids and these are sucked by means of their proboscis. They feed on nectar from flowers and also sip water from damp patches. This they do for water, for energy from sugars in nectar and for sodium and other minerals which are vital for their reproduction. Several species of butterflies need more sodium than provided by nectar. They are attracted to sodium in salt and they sometimes land on people, attracted by human sweat. Besides damp patches, some butterflies also visit dung, rotting fruit or carcasses to obtain minerals and nutrients. In many species, this Mud-puddling behaviour is restricted to the males and studies have suggested that the nutrients collected are provided as a nuptial gift along with the spermatophore during mating. Butterflies sense the air for scents, wind and nectar using their antennae. The antennae come in various shapes and colours. The hesperids have a pointed angle or hook to the antennae, while most other families show knobbed antennae. The antennae are richly covered with sensillae. A butterfly's sense of taste is coordinated by chemoreceptors on the tarsi, which work only on contact, and are used to determine whether an egg-laying insect's offspring will be able to feed on a leaf before eggs are laid on it. Many butterflies use chemical signals, pheromones, and specialized scent scales (androconia) and other structures (coremata or 'Hair pencils' in the Danaidae) are developed in some species. Vision is well developed in butterflies and most species are sensitive to the ultraviolet spectrum. Many species show sexual dimorphism in the patterns of UV reflective patches. Color vision may be widespread but has been demonstrated in only a few species. Some butterflies have organs of hearing and some species are also known to make stridulatory and clicking sounds. Many butterflies, such as the Monarch butterfly, are migratory and capable of long distance flights. They migrate during the day and use the sun to orient themselves. They also perceive polarized light and use it for orientation when the sun is hidden. Many species of butterfly maintain territories and actively chase other species or individuals that may stray into them. Some species will bask or perch on chosen perches. The flight styles of butterflies are often characteristic and some species have courtship flight displays. Basking is an activity which is more common in the cooler hours of the morning. Many species will orient themselves to gather heat from the sun. Some species have evolved dark wingbases to help in gathering more heat and this is especially evident in alpine forms. Flight. Like many other members of the insect world, the lift generated by butterflies is more than what can be accounted for by steady-state, non-transitory aerodynamics. Studies using "Vanessa atalanta" in a windtunnel show that they use a wide variety of aerodynamic mechanisms to generate force. These include wake capture, vortices at the wing edge, rotational mechanisms and Weis-Fogh 'clap-and-fling' mechanisms. The butterflies were also able to change from one mode to another rapidly. (See also Insect flight) Migration. Many butterflies migrate over long distances. Particularly famous migrations being those of the Monarch butterfly from Mexico to North America, a distance of about 4,000 to 4,800 kilometres (2500-3000 miles). Other well known migratory species include the Painted Lady and several of the Danaine butterflies. Spectacular and large scale migrations associated with the Monsoons are seen in peninsular India. Migrations have been studied in more recent times using wing tags and also using stable hydrogen isotopes. Butterflies have been shown to navigate using time compensated sun compasses. They can see polarized light and therefore orient even in cloudy conditions. The polarized light in the region close to the ultraviolet spectrum is suggested to be particularly important. It is suggested that most migratory butterflies are those that belong to semi-arid areas where breeding seasons are short. The life-histories of their host plants also influence the strategies of the butterflies. Defense. Butterflies are threatened in their early stages by parasitoids and in all stages by predators, diseases and environmental factors. They protect themselves by a variety of means. Chemical defenses are widespread and are mostly based on chemicals of plant origin. In many cases the plants themselves evolved these toxic substances as protection against herbivores. Butterflies have evolved mechanisms to sequester these plant toxins and use them instead in their own defense. These defense mechanisms are effective only if they are also well advertised and this has led to the evolution of bright colours in unpalatable butterflies. This signal may be mimicked by other butterflies. These mimetic forms are usually restricted to the females. Cryptic coloration is found in many butterflies. Some like the oakleaf butterfly are remarkable imitations of leaves. As caterpillars, many defend themselves by freezing and appearing like sticks or branches. Some papilionid caterpillars resemble bird dropping in their early instars. Some caterpillars have hairs and bristly structures that provide protection while others are gregarious and form dense aggregations. Some species also form associations with ants and gain their protection (See Myrmecophile). Behavioural defenses include perching and wing positions to avoid being conspicuous. Some female Nymphalid butterflies are known to guard their eggs from parasitoid wasps. Eyespots and tails are found in many lycaenid butterflies and these divert the attention of predators from the more vital head region. An alternative theory is that these cause ambush predators such as spiders to approach from the wrong end and allow for early visual detection. A butterfly's hind wings are thought to allow the butterfly to take, swift, tight turns to evade predators. Notable species. There are between 15,000 and 20,000 species of butterflies worldwide. Some well known species from around the world include: Art. Artistic depictions of butterflies have been used in many cultures including Egyptian hieroglyphs 3500 years ago. Today, butterflies are widely used in various objects of art and jewelry: mounted in frame, embedded in resin, displayed in bottles, laminated in paper, and used in some mixed media artworks and furnishings. Butterflies have also inspired the "butterfly fairy" as an art and fictional character. Symbolism. According to the “Butterflies” chapter in by Lafcadio Hearn, a butterfly is seen as the personification of a person's soul; whether they be living, dying, or already dead. One Japanese superstition says that if a butterfly enters your guestroom and perches behind the bamboo screen, the person whom you most love is coming to see you. However, large numbers of butterflies are viewed as bad omens. When Taira no Masakado was secretly preparing for his famous revolt, there appeared in Kyoto so vast a swarm of butterflies that the people were frightened -thinking the apparition to be a portent of coming evil. The Russian word for "butterfly", бабочка ("bábochka"), also means "bow tie". It is a diminutive of "baba" or "babka" ("woman, grandmother, cake", whence also "babushka"= "grandmother". The Ancient Greek word for "butterfly" is ψυχή ("psȳchē"), which primarily means "soul", "mind". According to Mircea Eliade's "Encyclopedia of Religion", some of the Nagas of Manipur trace their ancestry from a butterfly. In Chinese culture two butterflies flying together are a symbol of love. Also a famous Chinese folk story called Butterfly Lovers. The Taoist philosopher Zhuangzi once had a dream of being a butterfly flying without care about humanity, however when he woke up and realized it was just a dream, he thought to himself "Was I before a man who dreamt about being a butterfly, or am I now a butterfly who dreams about being a man?" In some old cultures, butterflies also symbolize rebirth into a new life after being inside a cocoon for a period of time. Jose Rizal delivered a speech in 1884 in a banquet and mentioned "the Oriental chrysalis... is about to leave its cocoon" comparing the emergence of a "new Philippines" with that of butterfly metamorphosis. He has also often used the butterfly imagery in his poems and other writings to express the Spanish Colonial Filipinos' longing for liberty. Much later, in a letter to Ferdinand Blumentritt, Rizal compared his life in exile to a weary butterfly with sun-burnt wings. Some people say that when a butterfly lands on you it means good luck. However, in Devonshire, people would traditionally rush around to kill the first butterfly of the year that they see, or else face a year of bad luck. Also, in the Philippines, a lingering black butterfly or moth in the house is taken to mean that someone in the family has died or will soon die. The idiom "butterflies in the stomach" is used to describe a state of nervousness. Technological inspiration. Researches on the wing structure of Palawan Birdwing butterflies led to new wide wingspan kite and aircraft designs. Studies on the reflection and scattering of light by the scales on wings of swallowtail butterflies led to the innovation of more efficient light-emitting diodes. The structural coloration of butterflies is inspiring nanotechnology research to produce paints that do not use toxic pigments and in the development of new display technologies. Furthermore, the discoloration and health of butterflies in butterfly farms, is now being studied for use as indicators of air quality in several cities.