Eye
Schematic diagram of the vertebrate eye.
Compound eye of Antarctic krill

Eyes are organs that detect light and convert it into electro-chemical impulses in neurons. The simplest photoreceptors in conscious vision connect light to movement. In higher organisms the eye is a complex optical system which collects light from the surrounding environment, regulates its intensity through a diaphragm, focuses it through an adjustable assembly of lenses to form an image, converts this image into a set of electrical signals, and transmits these signals to the brain through complex neural pathways that connect the eye via the optic nerve to the visual cortex and other areas of the brain. Eyes with resolving power have come in ten fundamentally different forms, and 96% of animal species possess a complex optical system.^[1] Image-resolving eyes are present in molluscs, chordates and arthropods.^[2]

The simplest "eyes", such as those in microorganisms, do nothing but detect whether the surroundings are light or dark, which is sufficient for the entrainment of circadian rhythms.^{[citation needed]} From more complex eyes, retinal photosensitive ganglion cells send signals along the retinohypothalamic tract to the suprachiasmatic nuclei to effect circadian adjustment.

Overview[link]

Eye of the wisent,
the European bison

Complex eyes can distinguish shapes and colours. 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, which have monocular vision.

The first proto-eyes evolved among animals 600 million years ago about the time of the Cambrian explosion.^[3] 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 six of the thirty-plus^[4] main phyla.^[1] In most vertebrates and some molluscs, the eye works by allowing light to enter and project onto a light-sensitive panel of cells, known as the retina, at the rear of the eye. The cone cells (for colour) 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,^[5] and reducing aberrations when there is enough light.^[6]

The eyes of most cephalopods, fish, amphibians and snakes have fixed lens shapes, and focusing vision is achieved by telescoping the lens—similar to how a camera focuses.^[7]

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° 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 colour vision, the Mantis shrimp has been reported to have the world's most complex colour vision system.^[8] 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.^[9]

Evolution[link]

Evolution of the eye

Photoreception is phylogenetically very old, with various theories of phylogenesis.^[10] The common origin (monophyly) of all animal eyes is now widely accepted as fact. This is based upon the shared anatomical and genetic features of all eyes; that is, all modern eyes, varied as they are, have their origins in a proto-eye believed to have evolved some 540 million years ago,^[11][12][13] and the PAX6 gene is considered a key factor in this. The majority of the advancements in early eyes are believed to have taken only a few million years to develop, since the first predator to gain true imaging would have touched off an "arms race".^[14] Prey animals and competing predators alike would be at a distinct disadvantage without such capabilities and would be less likely to survive and reproduce. Hence multiple eye types and subtypes developed in parallel.

Eyes in various animals show adaptation to their requirements. For example, birds of prey have much greater visual acuity than humans, and some can see ultraviolet light. The different forms of eye in, for example, vertebrates and molluscs are often cited as examples of parallel evolution, despite their distant common ancestry.

The very earliest "eyes", called eyespots, were simple patches of photoreceptor protein in unicellular animals. In multicellular beings, multicellular eyespots evolved, physically similar to the receptor patches for taste and smell. These eyespots could only sense ambient brightness: they could distinguish light and dark, but not the direction of the light source.^[1]

Through gradual change, as the eyespot depressed into a shallow "cup" shape, the ability to slightly discriminate directional brightness was achieved by using the angle at which the light hit certain cells to identify the source. The pit deepened over time, the opening diminished in size, and the number of photoreceptor cells increased, forming an effective pinhole camera that was capable of dimly distinguishing shapes.^[15]

The thin overgrowth of transparent cells over the eye's aperture, originally formed to prevent damage to the eyespot, allowed the segregated contents of the eye chamber to specialise into a transparent humour that optimised colour filtering, blocked harmful radiation, improved the eye's refractive index, and allowed functionality outside of water. The transparent protective cells eventually split into two layers, with circulatory fluid in between that allowed wider viewing angles and greater imaging resolution, and the thickness of the transparent layer gradually increased, in most species with the transparent crystallin protein.^[16]

The gap between tissue layers naturally formed a bioconvex shape, an optimally ideal structure for a normal refractive index. Independently, a transparent layer and a nontransparent layer split forward from the lens: the cornea and iris. Separation of the forward layer again formed a humour, the aqueous humour. This increased refractive power and again eased circulatory problems. Formation of a nontransparent ring allowed more blood vessels, more circulation, and larger eye sizes.^[16]

Types of eye[link]

There are ten different eye layouts—indeed every way of capturing an optical image commonly used by human beings, with the exceptions of zoom and Fresnel lenses. Eye types can be categorised into "simple eyes", with one concave photoreceptive surface, and "compound eyes", which comprise a number of individual lenses laid out on a convex surface.^[1] 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.^[1] 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 rhabdomeric. These two groups are not monophyletic; the cnidaria also possess cilliated cells, ^[17] and some annelids possess both.^[18]

Non-compound eyes[link]

Simple eyes are rather ubiquitous, and lens-bearing eyes have evolved at least seven times in vertebrates, cephalopods, annelids, crustaceans and cubozoa.^[19]

Pit eyes[link]

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.^[1] 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.^[1] 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.^[1]

Pit vipers have developed pits that function as eyes by sensing thermal infra-red radiation, in addition to their optical wavelength eyes like those of other vertebrates.

Spherical lensed eye[link]

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.^[1] The most basic form, 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.^[1] 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.^[1] 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.^[1]

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.^[1] No aquatic organisms possess homogeneous lenses; presumably the evolutionary pressure for a heterogeneous lens is great enough for this stage to be quickly "outgrown".^[1]

This eye creates an image that is sharp enough that motion of the eye can cause significant blurring. To minimise the effect of eye motion while the animal moves, most such eyes have stabilising eye muscles.^[1]

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.^[20] Focusing the image would also cause the sun's image to be focused 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.^[20] 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).^[20]

Multiple lenses[link]

Some marine organisms bear more than one lens; for instance the copepod Pontella has three. The outer has a parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed. Another copepod, Copilia, has two lenses in each eye, arranged like those in a telescope.^[1] 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.^[1]

Refractive cornea[link]

In the eyes of most mammals, birds, reptiles, and most other terrestrial vertebrates (along with spiders and some insect larvae) the vitreous fluid has a higher refractive index than the air.^[1] In general, the lens is not spherical. Spherical lenses produce spherical aberration. In refractive corneas, the lens tissue is corrected with inhomogeneous lens material (see Luneburg lens), or with an aspheric shape.^[1] Flattening the lens has a disadvantage; the quality of vision is diminished away from the main line of focus. Thus, animals that have evolved with a wide field-of-view often have eyes that make use of an inhomogeneous lens.^[1]

As mentioned above, a refractive cornea is only useful out of water; in water, there is little difference in refractive index between the vitreous fluid and the surrounding water. Hence creatures that have returned to the water–penguins and seals, for example–lose their highly curved cornea and return to lens-based vision. An alternative solution, borne by some divers, is to have a very strongly focusing cornea.^[1]

Reflector eyes[link]

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.^[1] 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.^[1]

Many small organisms such as rotifers, copepods and platyhelminths use such organs, but these are too small to produce usable images.^[1] Some larger organisms, such as scallops, also use reflector eyes. The scallop Pecten has up to 100 millimetre-scale reflector eyes fringing the edge of its shell. It detects moving objects as they pass successive lenses.^[1]

There is at least one vertebrate, the spookfish, whose eyes include reflective optics for focusing of light. Each of the two eyes of a spookfish collects light from both above and below; the light coming from above is focused by a lens, while that coming from below, by a curved mirror composed of many layers of small reflective plates made of guanine crystals.^[21]

[edit] Compound eyes

An image of a house fly compound eye surface by using scanning electron microscope

Anatomy of the compound eye of an insect

Arthropods such as this Calliphora vomitoria fly have compound eyes

A compound eye may consist of thousands of individual photoreceptor units or ommatidia (ommatidium, singular). 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 polarisation of light.^[22] Because the individual lenses are so small, the effects of diffraction impose a limit on the possible resolution that can be obtained (assuming that they do not function as phased arrays). 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 heads.^{[citation needed]}

Compound eyes fall into two groups: apposition eyes, which form multiple inverted images, and superposition eyes, which form a single erect image.^[23] Compound eyes are common in arthropods, and are also present in annelids and some bivalved molluscs.^[24]

Compound eyes, in arthropods at least, grow at their margins by the addition of new ommatidia.^[25]

Apposition eyes[link]

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.^[1] 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.^[1] (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.^[1]

Superposition eyes[link]

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 have a transparent gap but use corner mirrors instead of lenses.

Parabolic superposition[link]

This eye type functions by refracting light, then using a parabolic mirror to focus the image; it combines features of superposition and apposition eyes.^[9]

Other[link]

Good fliers like flies or honey bees, or prey-catching insects like praying mantis or dragonflies, have specialised zones of ommatidia organised into a fovea area which gives acute vision. In the acute zone the eyes 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 specialised 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, organised 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. The tube feet of sea urchins contain photoreceptor proteins, which together act as a compound eye; they lack screening pigments, but can detect the directionality of light by the shadow cast by its opaque body.^[26]

Nutrients[link]

The ciliary body is triangular in horizontal section and is coated by a double layer, the ciliary epithelium. The inner layer is transparent and covers the vitreous body, and is continuous from the neural tissue of the retina. The outer layer is highly pigmented, continuous with the retinal pigment epithelium, and constitutes the cells of the dilator muscle.

The vitreous is the transparent, colourless, gelatinous mass that fills the space between the lens of the eye and the retina lining the back of the eye.^[27] It is produced by certain retinal cells. It is of rather similar composition to the cornea, but contains very few cells (mostly phagocytes which remove unwanted cellular debris in the visual field, as well as the hyalocytes of Balazs of the surface of the vitreous, which reprocess the hyaluronic acid), no blood vessels, and 98-99% of its volume is water (as opposed to 75% in the cornea) with salts, sugars, vitrosin (a type of collagen), a network of collagen type II fibres with the mucopolysaccharide hyaluronic acid, and also a wide array of proteins in micro amounts. Amazingly, with so little solid matter, it tautly holds the eye.

Relationship to life requirements[link]

Eyes are generally adapted to the environment and life requirements 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.^[1]:23-4 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.^[28] 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.^[29] 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.^[28] 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.^[28] 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.^[9]

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.^[28] 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.^[28]

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.^[28]

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.^[9]

Visual acuity[link]

The eye of a red-tailed hawk

Visual acuity, or resolving power, is "the ability to distinguish fine detail" and is the property of cone cells.^[30] It 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 grey block, is then the measurement of visual acuity of the eye.

For a human eye with excellent acuity, the maximum theoretical resolution is 50 CPD^[31] (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.^[32] 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.^{[citation needed]}

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.^[33] 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.

However, in the compound eye, the resolution is related to the size of individual ommatidia and the distance between neighbouring ommatidia. Physically these cannot be reduced in size to achieve the acuity seen with single lensed eyes as in mammals. Compound eyes have a much lower acuity than mammalian eyes.^[34]

Perception of colors[link]

"Color vision is the faculty of the organism to distinguish lights of different spectral qualities."^[35] All organisms are restricted to a small range of electromagnetic spectrum; this varies from creature to creature, but is mainly between wavelengths of 400 and 700 nm.^[36] This is a rather small section of the electromagnetic spectrum, probably reflecting the submarine evolution of the organ: water blocks out all but two small windows of the EM spectrum, and there has been no evolutionary pressure among land animals to broaden this range.^[37]

The most sensitive pigment, rhodopsin, has a peak response at 500 nm.^[38] Small changes to the genes coding for this protein can tweak the peak response by a few nm;^[2] pigments in the lens can also filter incoming light, changing the peak response.^[2] Many organisms are unable to discriminate between colours, seeing instead in shades of grey; colour vision necessitates a range of pigment cells which are primarily sensitive to smaller ranges of the spectrum. In primates, geckos, and other organisms, these take the form of cone cells, from which the more sensitive rod cells evolved.^[38] Even if organisms are physically capable of discriminating different colours, this does not necessarily mean that they can perceive the different colours; only with behavioural tests can this be deduced.^[2]

Most organisms with color vision are able to detect ultraviolet light. This high energy light can be damaging to receptor cells. With a few exceptions (snakes, placental mammals), most organisms avoid these effects by having absorbent oil droplets around their cone cells. The alternative, developed by organisms that had lost these oil droplets in the course of evolution, is to make the lens impervious to UV light — this precludes the possibility of any UV light being detected, as it does not even reach the retina.^[38]

Rods and cones[link]

The retina contains two major types of light-sensitive photoreceptor cells used for vision: the rods and the cones.

Rods cannot distinguish colours, but are responsible for low-light (scotopic) monochrome (black-and-white) vision; they work well in dim light as they contain a pigment, rhodopsin (visual purple), which is sensitive at low light intensity, but saturates at higher (photopic) intensities. Rods are distributed throughout the retina but there are none at the fovea and none at the blind spot. Rod density is greater in the peripheral retina than in the central retina.

Cones are responsible for colour vision. They require brighter light to function than rods require. In humans, there are three types of cones, maximally sensitive to long-wavelength, medium-wavelength, and short-wavelength light (often referred to as red, green, and blue, respectively, though the sensitivity peaks are not actually at these colours). The colour seen is the combined effect of stimuli to, and responses from, these three types of cone cells. Cones are mostly concentrated in and near the fovea. Only a few are present at the sides of the retina. Objects are seen most sharply in focus when their images fall on the fovea, as when one looks at an object directly. Cone cells and rods are connected through intermediate cells in the retina to nerve fibres of the optic nerve. When rods and cones are stimulated by light, the nerves send off impulses through these fibres to the brain.^[38]

Pigmentation[link]

The pigment molecules used in the eye are various, but can be used to define the evolutionary distance between different groups, and can also be an aid in determining which are closely related – although problems of convergence do exist.^[38]

Opsins are the pigments involved in photoreception. Other pigments, such as melanin, are used to shield the photoreceptor cells from light leaking in from the sides. The opsin protein group evolved long before the last common ancestor of animals, and has continued to diversify since.^[2]

There are two types of opsin involved in vision; c-opsins, which are associated with ciliary-type photoreceptor cells, and r-opsins, associated with rhabdomeric photoreceptor cells.^[39] The eyes of vertebrates usually contain cilliary cells with c-opsins, and (bilaterian) invertebrates have rhabdomeric cells in the eye with r-opsins. However, some ganglion cells of vertebrates express r-opsins, suggesting that their ancestors used this pigment in vision, and that remnants survive in the eyes.^[39] Likewise, c-opsins have been found to be expressed in the brain of some invertebrates. They may have been expressed in ciliary cells of larval eyes, which were subsequently resorbed into the brain on metamorphosis to the adult form.^[39] C-opsins are also found in some derived bilaterian-invertebrate eyes, such as the pallial eyes of the bivalve molluscs; however, the lateral eyes (which were presumably the ancestral type for this group, if eyes evolved once there) always use r-opsins.^[39] Cnidaria, which are an outgroup to the taxa mentioned above, express c-opsins - but r-opsins are yet to be found in this group.^[39] Incidentally, the melanin produced in the raticate is produced in the same fashion as that in vertebrates, suggesting the common descent of this pigment.^[39]

See also[link]

	Book: Eye
Wikipedia books are collections of articles that can be downloaded or ordered in print.

• Arthropod eye
• Cephalopod eye
• Emission theory (vision)
• Eye colour
• Eyelid
• Eye movement (sensory)
• Human eye
• Mammalian eye
• Mollusc eye
• Nictitating membrane
• Simple eye in invertebrates
• Tapetum lucidum
• Tears

References[link]

Notes[link]

Bibliography[link]

• Ali, Mohamed Ather; Klyne, M. A. (1985). Vision in Vertebrates. New York: Plenum Press. ISBN 0-306-42065-1. 

External links[link]

Wikimedia Commons has media related to: Eyes

• Evolution of the eye
• Diagram of the eye
• Webvision. The organisation of the retina and visual system. An in-depth treatment of retinal function, open to all but geared most toward graduate students.
• Eye strips images of all but bare essentials before sending visual information to brain, UC Berkeley research shows
• A collection of resources for eye anatomy.
• Anatomy of the eye - flash animated interactive.
• Eyes - Eyes are organs that detect light, and send signals along the optic nerve to the visual and other areas of the brain.
• Detection of weak optical signals by the human visual system : Perspectives in Neuroscience and in Quantum Physics
• Are Your Eyes Right February 1944 Popular Science

v t e Sensory system – visual system – globe of eye (TA A15.2.1–6, TH 3.11.08.0-5, GA 10.1005) Fibrous tunic (outer) Sclera Episcleral layer • Schlemm's canal • Trabecular meshwork Cornea Limbus • layers (Epithelium, Bowman's, Stroma, Descemet's, Endothelium) Uvea/vascular tunic (middle) Choroid Capillary lamina of choroid • Bruch's membrane • Sattler's layer Ciliary body Ciliary processes • Ciliary muscle Iris Stroma • Pupil • Iris dilator muscle • Iris sphincter muscle Retina (inner) Layers Inner limiting membrane • Nerve fiber layer • Ganglion cell layer • Inner plexiform layer • Inner nuclear layer Outer plexiform layer • Outer nuclear layer External limiting membrane • Layer of rods and cones • Retinal pigment epithelium Cells Photoreceptor cells (Cone cell, Rod cell) → (Horizontal cell) → Bipolar cell → (Amacrine cell) → Retina ganglion cell (Midget cell, Parasol cell, Bistratified cell, Giant retina ganglion cells, Photosensitive ganglion cell) → Diencephalon: P cell, M cell, K cell Muller glia Other Macula (Foveola, Fovea centralis) • Optic disc (Optic cup) Anterior segment Anterior chamber • Aqueous humour • Posterior chamber • Lens (Capsule of lens, Zonule of Zinn) Posterior segment Vitreous humour Other Ocular immune system • Tapetum lucidum • Keratocytes M: EYE anat(g/a/p)/phys/devp/prot noco/cong/tumr, epon proc, drug(S1A/1E/1F/1L)

Bette Davis
Studio publicity photo, c. 1935
Born	Ruth Elizabeth Davis (1908-04-05)April 5, 1908 Lowell, Massachusetts, U.S.
Died	October 6, 1989(1989-10-06) (aged 81) Neuilly-sur-Seine, France
Cause of death	Breast cancer
Occupation	Actress
Years active	1929–1989
Spouse	Harmon Nelson (m.1932-1938; divorced) Arthur Farnsworth (m.1940-1943; his death) William Grant Sherry (m.1945-1950; divorced) Gary Merrill (m.1950-1960; divorced)
Children	Barbara Merrill, born on (1947-05-01) May 1, 1947 (age 65) Michael Merrill Margot Merrill

Ruth Elizabeth "Bette" Davis (April 5, 1908 – October 6, 1989) was an American actress of film, television and theater. Noted for her willingness to play unsympathetic characters, she was highly regarded for her performances in a range of film genres, from contemporary crime melodramas to historical and period films and occasional comedies, although her greatest successes were her roles in romantic dramas.

After appearing in Broadway plays, Davis moved to Hollywood in 1930, but her early films for Universal Studios were unsuccessful. She joined Warner Bros. in 1932 and established her career with several critically acclaimed performances. In 1937, she attempted to free herself from her contract and although she lost a well-publicized legal case, it marked the beginning of the most successful period of her career. Until the late 1940s, she was one of American cinema's most celebrated leading ladies, known for her forceful and intense style. Davis gained a reputation as a perfectionist who could be highly combative, and confrontations with studio executives, film directors and costars were often reported. Her forthright manner, clipped vocal style and ubiquitous cigarette contributed to a public persona which has often been imitated and satirized.

Davis was the co-founder of the Hollywood Canteen, and was the first female president of the Academy of Motion Picture Arts and Sciences. She won the Academy Award for Best Actress twice, was the first person to accrue 10 Academy Award nominations for acting, and was the first woman to receive a Lifetime Achievement Award from the American Film Institute. Her career went through several periods of eclipse, and she admitted that her success had often been at the expense of her personal relationships. Married four times, she was once widowed and thrice divorced, and raised her children as a single parent. Her final years were marred by a long period of ill health, but she continued acting until shortly before her death from breast cancer, with more than 100 films, television and theater roles to her credit. In 1999, Davis was placed second, after Katharine Hepburn, on the American Film Institute's list of the greatest female stars of all time.

Life and career[link]

Background and early acting career (1908–1929)[link]

Ruth Elizabeth Davis, known from early childhood as "Betty", was born in Lowell, Massachusetts, the daughter of Ruth Augusta "Ruthie" (née Favor) and Harlow Morrell Davis, a patent attorney;^[1] her sister, Barbara "Bobby", was born October 25, 1909. The family was Protestant, of English, French, and Welsh ancestry. In 1915, Davis's parents separated and Betty and Bobby attended a Spartan boarding school called Crestalban in Lanesborough, which is located in the Berkshires.^[2] In 1921, Ruth Davis moved to New York City with her daughters, where she worked as a portrait photographer. Betty was inspired to become an actress after seeing Rudolph Valentino in The Four Horsemen of the Apocalypse (1921) and Mary Pickford in Little Lord Fauntleroy (1921),^[3] and changed the spelling of her name to "Bette" after Honoré de Balzac's La Cousine Bette.^[4] She received encouragement from her mother, who had aspired to become an actress.

She attended Cushing Academy, a boarding school in Ashburnham, Massachusetts, where she met her future husband, Harmon O. Nelson, known as "Ham". In 1926, she saw a production of Henrik Ibsen's The Wild Duck with Blanche Yurka and Peg Entwistle. Davis later recalled that it inspired her full commitment to her chosen career, and said, "Before that performance I wanted to be an actress. When it ended, I had to be an actress ... exactly like Peg Entwistle."^[5] She auditioned for admission to Eva LeGallienne's Manhattan Civic Repertory, but was rejected by LeGallienne who described her attitude as "insincere" and "frivolous".^[6] She was accepted by the John Murray Anderson School of Theatre, and studied dance with Martha Graham.

She auditioned for George Cukor's stock theater company, and although he was not very impressed, he gave Davis her first paid acting assignment anyway—a one-week stint playing the part of a chorus girl in the play Broadway. She was later chosen to play Hedwig, the character she had seen Entwistle play, in The Wild Duck. After performing in Philadelphia, Washington and Boston, she made her Broadway debut in 1929 in Broken Dishes, and followed it with Solid South. A Universal Studios talent scout saw her perform and invited her to Hollywood for a screen test.

Early years in Hollywood (1930–1936)[link]

Davis in her film debut, The Bad Sister (1931)

Accompanied by her mother, Davis traveled by train to Hollywood, arriving on December 13, 1930. She later recounted her surprise that nobody from the studio was there to meet her; a studio employee had waited for her, but left because he saw nobody who "looked like an actress". She failed her first screen test but was used in several screen tests for other actors. In a 1971 interview with Dick Cavett, she related the experience with the observation, "I was the most Yankee-est, most modest virgin who ever walked the earth. They laid me on a couch, and I tested fifteen men ... They all had to lie on top of me and give me a passionate kiss. Oh, I thought I would die. Just thought I would die."^[7] A second test was arranged for Davis, for the film A House Divided (1931). Hastily dressed in an ill-fitting costume with a low neckline, she was rebuffed by the director William Wyler, who loudly commented to the assembled crew, "What do you think of these dames who show their chests and think they can get jobs?"^[8] Carl Laemmle, the head of Universal Studios, considered terminating Davis's employment, but cinematographer Karl Freund told him she had "lovely eyes" and would be suitable for The Bad Sister (1931), in which she subsequently made her film debut.^[9] Her nervousness was compounded when she overheard the Chief of Production, Carl Laemmle, Jr., comment to another executive that she had "about as much sex appeal as Slim Summerville", one of the film's co-stars.^[10] The film was not a success, and her next role in Seed (1931) was too brief to attract attention.

Universal Studios renewed her contract for three months, and she appeared in a small role in Waterloo Bridge (1931) before being lent to Columbia Pictures for The Menace and to Capital Films for Hell's House (all 1932). After nine months, and six unsuccessful films, Laemmle elected not to renew her contract. George Arliss chose Davis for the lead female role in The Man Who Played God (1932), and for the rest of her life, Davis credited him with helping her achieve her "break" in Hollywood. The Saturday Evening Post wrote, "she is not only beautiful, but she bubbles with charm", and compared her to Constance Bennett and Olive Borden.^[11] Warner Bros. signed her to a five-year contract.

In 1932, she married Harmon "Ham" Nelson, who was scrutinized by the press; his $100 a week earnings compared unfavorably with Davis's reported $1,000 a week income. Davis addressed the issue in an interview, pointing out that many Hollywood wives earned more than their husbands, but the situation proved difficult for Nelson, who refused to allow Davis to purchase a house until he could afford to pay for it himself.^[12] Davis had several abortions during the marriage.^[13]

As the shrewish Mildred in Of Human Bondage (1934), Davis was acclaimed for her dramatic performance.

After more than 20 film roles, the role of the vicious and slatternly Mildred Rogers in the RKO Radio production of Of Human Bondage (1934), a film adaptation of W. Somerset Maugham's novel, earned Davis her first major critical acclaim. Many actresses feared playing unsympathetic characters and several had refused the role, but Davis viewed it as an opportunity to show the range of her acting skills. Her costar, Leslie Howard, was initially dismissive of her, but as filming progressed his attitude changed and he subsequently spoke highly of her abilities. The director, John Cromwell, allowed her relative freedom, and commented, "I let Bette have her head. I trusted her instincts." She insisted that she be portrayed realistically in her death scene, and said, "the last stages of consumption, poverty and neglect are not pretty and I intended to be convincing-looking".^[14]

The film was a success, and Davis's confronting characterization won praise from critics, with Life magazine writing that she gave "probably the best performance ever recorded on the screen by a U.S. actress".^[15] Davis anticipated that her reception would encourage Warner Bros. to cast her in more important roles, and was disappointed when Jack Warner refused to lend her to Columbia Studios to appear in It Happened One Night, and instead cast her in the melodrama Housewife.^[16] When Davis was not nominated for an Academy Award for Of Human Bondage, The Hollywood Citizen News questioned the omission and Norma Shearer, herself a nominee, joined a campaign to have Davis nominated. This prompted an announcement from the Academy president, Howard Estabrook, who said that under the circumstances "any voter ... may write on the ballot his or her personal choice for the winners", thus allowing, for the only time in the Academy's history, the consideration of a candidate not officially nominated for an award.^[17] Claudette Colbert won the award for It Happened One Night but the uproar led to a change in Academy voting procedures the following year, whereby nominations were determined by votes from all eligible members of a particular branch, rather than by a smaller committee,^[18] with results independently tabulated by the accounting firm Price Waterhouse.^[19]

Davis appeared in Dangerous (1935) as a troubled actress and received very good reviews. E. Arnot Robertson wrote in Picture Post, "I think Bette Davis would probably have been burned as a witch if she had lived two or three hundred years ago. She gives the curious feeling of being charged with power which can find no ordinary outlet." The New York Times hailed her as "becoming one of the most interesting of our screen actresses".^[20] She won the Academy Award for Best Actress for the role, but commented it was belated recognition for Of Human Bondage, calling the award a "consolation prize".^[21] For the rest of her life, Davis maintained that she gave the statue its familiar name of "Oscar" because its posterior resembled that of her husband, whose middle name was Oscar,^[22][23] although her claim has been disputed by the Academy of Motion Picture Arts and Sciences, among others.

In her next film, The Petrified Forest (1936), Davis co-starred with Leslie Howard and Humphrey Bogart, but Bogart, in his first important role, received most of the critics' praise. Davis appeared in several films over the next two years but most were poorly received.

Legal case[link]

Convinced that her career was being damaged by a succession of mediocre films, Davis accepted an offer in 1936 to appear in two films in England. Knowing that she was breaching her contract with Warner Bros., she fled to Canada to avoid legal papers being served upon her. Eventually, Davis brought her case to court in England, hoping to get out of her contract with Warner Bros.^[24] She later recalled the opening statement of the barrister, Sir Patrick Hastings, who represented Warner Bros. Hastings urged the court to "come to the conclusion that this is rather a naughty young lady and that what she wants is more money". He mocked Davis's description of her contract as "slavery" by stating, incorrectly, that she was being paid $1,350 per week. He remarked, "if anybody wants to put me into perpetual servitude on the basis of that remuneration, I shall prepare to consider it." The British press offered little support to Davis, and portrayed her as overpaid and ungrateful.^[25]

Davis explained her viewpoint to a journalist, saying "I knew that, if I continued to appear in any more mediocre pictures, I would have no career left worth fighting for."^[26] Davis's counsel presented her complaints—that she could be suspended without pay for refusing a part, with the period of suspension added to her contract, that she could be called upon to play any part within her abilities regardless of her personal beliefs, that she could be required to support a political party against her beliefs, and that her image and likeness could be displayed in any manner deemed applicable by the studio. Jack Warner testified, and was asked, "Whatever part you choose to call upon her to play, if she thinks she can play it, whether it is distasteful and cheap, she has to play it?" Warner replied, "Yes, she must play it."^[27] The case, decided by Branson J. in the English High Court, was reported as Warner Bros. Studios Incorporated v. Nelson in [1937] 1 KB 209. Davis lost the case and returned to Hollywood, in debt and without income, to resume her career. Olivia de Havilland mounted a similar case in 1943 and won.

Success with Warner Bros. (1937–1941)[link]

Jezebel (1938) was Davis's second Academy Award-winning performance

Davis began work on Marked Woman (1937), as a prostitute in a contemporary gangster drama inspired by the case of Lucky Luciano. For her performance in the film she was awarded the Volpi Cup at the 1937 Venice Film Festival.^[28] Her next picture was Jezebel (1938), and during production Davis entered a relationship with director William Wyler. She later described him as the "love of my life", and said that making the film with him was "the time in my life of my most perfect happiness".^[29] The film was a success, and Davis's performance as a spoiled Southern belle earned her a second Academy Award, which led to speculation in the press that she would be chosen to play a similar character, Scarlett O'Hara, in Gone with the Wind. Davis expressed her desire to play Scarlett, and while David O. Selznick was conducting a search for the actress to play the role, a radio poll named her as the audience favorite. Warner offered her services to Selznick as part of a deal that also included Errol Flynn and Olivia de Havilland, but Selznick did not consider Davis as suitable, and rejected the offer,^[30] while Davis did not want Flynn cast as Rhett Butler.

Jezebel marked the beginning of the most successful phase of Davis's career, and over the next few years she was listed in the annual "Quigley Poll of the Top Ten Money Making Stars", which was compiled from the votes of movie exhibitors throughout the U.S. for the stars that had generated the most revenue in their theaters over the previous year.^[31] In contrast to Davis's success, her husband, Ham Nelson, had failed to establish a career for himself, and their relationship faltered. In 1938, Nelson obtained evidence that Davis was engaged in a sexual relationship with Howard Hughes and subsequently filed for divorce citing Davis's "cruel and inhuman manner".^[32]

She was emotional during the making of her next film, Dark Victory (1939), and considered abandoning it until the producer Hal B. Wallis convinced her to channel her despair into her acting. The film became one of the highest grossing films of the year, and the role of Judith Traherne brought her an Academy Award nomination. In later years, Davis cited this performance as her personal favorite.^[33] She appeared in three other box office hits in 1939, The Old Maid with Miriam Hopkins, Juarez with Paul Muni and The Private Lives of Elizabeth and Essex with Errol Flynn. The latter was her first color film and her only color film made during the height of her career. To play the elderly Elizabeth I of England, Davis shaved her hairline and eyebrows. During filming she was visited on the set by the actor Charles Laughton. She commented that she had a "nerve" playing a woman in her sixties, to which Laughton replied, "Never not dare to hang yourself. That's the only way you grow in your profession. You must continually attempt things that you think are beyond you, or you get into a complete rut." Recalling the episode many years later, Davis remarked that Laughton's advice had influenced her throughout her career.^[34]

Davis often played unlikable characters, such as Regina Giddens in The Little Foxes (1941).

By this time, Davis was Warner Bros.' most profitable star, and she was given the most important of their female leading roles. Her image was considered with more care; although she continued to play character roles, she was often filmed in close-ups that emphasized her distinctive eyes. All This and Heaven Too (1940) was the most financially successful film of Davis's career to that point, while The Letter (1940) was considered "one of the best pictures of the year" by The Hollywood Reporter, and Davis won admiration for her portrayal of an adulterous killer, a role originated by famed actress Katharine Cornell.^[35] During this time, she was in a relationship with her former costar George Brent, who proposed marriage. Davis refused, as she had met Arthur Farnsworth, a New England innkeeper. Davis and Farnsworth were married at Home Ranch, in Lake Montezuma, Arizona, in December 1940.

In January 1941, Davis became the first female president of the Academy of Motion Picture Arts and Sciences but antagonized the committee members with her brash manner and radical proposals. Faced with the disapproval and resistance of the committee, Davis resigned, and was succeeded by Jean Hersholt, who implemented the changes she had suggested. Davis starred in three movies in 1941, the first being The Great Lie, opposite George Brent. It was a refreshingly different role for Davis, as she played a kind, sympathetic character. Brent tickled Davis during many of the film's scenes, which allowed the audience, used to Davis' strong-willed character, a rare glimpse of her succumbing to giggles and squirms. William Wyler directed Davis for the third time in Lillian Hellman's The Little Foxes (RKO, 1941), but they clashed over the character of Regina Giddens. Taking a role originally played on stage by Tallulah Bankhead, Davis felt Bankhead's original interpretation was appropriate and followed Hellman's intent, but Wyler wanted her to soften the character. Davis refused to compromise. She received another Academy Award nomination for her performance, and never worked with Wyler again.

War effort and personal tragedy (1942–1944)[link]

Following the attack on Pearl Harbor, Davis spent the early months of 1942 selling war bonds. After Jack Warner criticized her tendency to cajole crowds into buying, she reminded him that her audiences responded most strongly to her "bitch" performances. She sold two million dollars worth of bonds in two days, as well as a picture of herself in Jezebel for $250,000. She also performed for black regiments as the only white member of an acting troupe formed by Hattie McDaniel, which included Lena Horne and Ethel Waters.^[36]

Paul Henreid lights Davis's cigarette in Now, Voyager (1942): an iconic scene

At John Garfield's suggestion of opening a servicemen's club in Hollywood, Davis—with the aid of Warner, Cary Grant and Jule Styne—transformed an old nightclub into the Hollywood Canteen, which opened on October 3, 1942. Hollywood's most important stars volunteered to entertain servicemen. Davis ensured that every night there would be a few important "names" for the visiting soldiers to meet.^[37] She appeared as herself in the film Hollywood Canteen (1944), which used the canteen as the setting for a fictional story. Davis later commented, "There are few accomplishments in my life that I am sincerely proud of. The Hollywood Canteen is one of them." In 1980, she was awarded the Distinguished Civilian Service Medal, the United States Department of Defense's highest civilian award, for her work with the Hollywood Canteen.^[38]

Davis showed little interest in the film Now, Voyager (1942) until Hal Wallis advised her that female audiences needed romantic dramas to distract them from the reality of their lives. It became one of the best known of her "women's pictures." In one of the film's most imitated scenes, Paul Henreid lights two cigarettes as he stares into Davis's eyes and passes one to her. Film reviewers complimented Davis on her performance, the National Board of Review commenting that she gave the film "a dignity not fully warranted by the script".^[39]

During the early 1940s, several of Davis's film choices were influenced by the war, such as Watch on the Rhine (1943), by Lillian Hellman, and Thank Your Lucky Stars (1943), a lighthearted all-star musical cavalcade, with each of the featured stars donating their fee to the Hollywood Canteen. Davis performed a novelty song, "They're Either Too Young or Too Old", which became a hit record after the film's release. Old Acquaintance (1943) reunited her with Miriam Hopkins in a story of two old friends who deal with the tensions created when one of them becomes a successful novelist. Davis felt that Hopkins tried to upstage her throughout the film. Director Vincent Sherman recalled the intense competitiveness and animosity between the two actresses, and Davis often joked that she held back nothing in a scene in which she was required to shake Hopkins in a fit of anger.^[40]

In August 1943, Davis's husband, Arthur Farnsworth, collapsed while walking along a Hollywood street and died two days later. An autopsy revealed that his fall had been caused by a skull fracture he had suffered two weeks earlier. Davis testified before an inquest that she knew of no event that might have caused the injury. A finding of accidental death was reached. Highly distraught, Davis attempted to withdraw from her next film Mr. Skeffington (1944); but Jack Warner, who had halted production following Farnsworth's death, convinced her to continue. Although she had gained a reputation for being forthright and demanding, her behavior during filming of Mr. Skeffington was erratic and out of character. She alienated Vincent Sherman by refusing to film certain scenes and insisting that some sets be rebuilt. She improvised dialogue, causing confusion among other actors, and infuriated the writer, Julius Epstein, who was called upon to rewrite scenes at her whim. Davis later explained her actions with the observation, "when I was most unhappy I lashed out rather than whined". Some reviewers criticized Davis for the excess of her performance; James Agee wrote that she "demonstrates the horrors of egocentricity on a marathonic scale";^[41] but despite the mixed reviews, she received another Academy Award nomination.

Professional setbacks (1945–1949)[link]

In The Corn Is Green (1945). Despite the studio's suggestion that Davis play the role as a young woman, the 38-year-old insisted on aging her appearance to fit the part.

In 1945, Davis married artist William Grant Sherry, who also, when necessary, worked as a masseur. She had been drawn to him because he claimed he had never heard of her and was, therefore, not intimidated by her.^[42] The same year, Davis refused the title role in Mildred Pierce (1945),^[43] a role for which Joan Crawford won an Academy Award, and instead made The Corn Is Green (1945) based on a play by Emlyn Williams. Davis played Miss Moffat, an English teacher who saves a young Welsh miner (John Dall) from a life in the coal pits, by offering him education. The part had been played in the theatre by Ethel Barrymore, but Warner Bros. felt that the film version should depict the character as a younger woman. Davis disagreed and insisted on playing the part as written and wore a gray wig and padding under her clothes, to create a dowdy appearance.^[44] The film was well received by critics and made a profit of $2.2 million.^[45] The critic E. Arnot Robertson observed that "only Bette Davis . . . could have combated so successfully the obvious intention of the adaptors of the play to make frustrated sex the mainspring of the chief character's interest in the young miner." He concluded that "the subtle interpretation she insisted on giving" kept the focus on the teacher's "sheer joy in imparting knowledge".^[46]

Her next film, A Stolen Life (1946), was the first and only film that Davis made with her own production company, BD Productions.^[47] Davis played dual roles, as twins. The film received poor reviews and was described by Bosley Crowther as "a distressingly empty piece";^[48] but, with a profit of $2.5 million, it was one of her biggest box-office successes.^[49] In 1947, the U.S. Treasury named Davis as the highest paid woman in the country,^[50] with her share of the film's profit accounting for most of her earnings. Her next film was Deception (1946), the first of her films to lose money.^[51]

Possessed (1947) had been tailor-made for Davis^[52] and was to have been her next project after Deception. However, she was pregnant and went on maternity leave. Joan Crawford played her role in Possessed and was nominated for an Academy Award as Best Actress. In 1947, at the age of 39, Davis gave birth to a daughter, Barbara Davis Sherry (known as B.D.) and later wrote in her memoir that she became absorbed in motherhood and considered ending her career. However, her relationship with Sherry began to deteriorate and she continued making films, but her popularity with audiences was steadily declining.^[53]

Among the film roles offered to Davis following her return to film making was Rose Sayer in The African Queen (1951). When informed that the film was to be made in Africa, Davis refused the part, telling Jack Warner, "If you can't shoot the picture in a boat on the back lot, then I'm not interested." Katharine Hepburn played the role.^[54] Davis was also offered a role in a film version of the Virginia Kellogg prison drama Women Without Men. Originally intended to pair Davis with Joan Crawford, Davis made it clear that she would not appear in any "dyke movie". It was filmed as Caged (1950), and the lead roles were played by Eleanor Parker and Agnes Moorehead.^[55] She lobbied Jack Warner to make two films, Ethan Frome and a biography of Mary Todd Lincoln; however, Warner vetoed each proposal.

Beyond the Forest (1949) was the last film Davis made for Warner Bros., after 17 years with the studio

In 1948, Davis was cast in the melodrama Winter Meeting; and, although she was initially enthusiastic, she soon learned that Warner had arranged for "softer" lighting to be used to disguise her age. She recalled that she had seen the same lighting technique "on the sets of Ruth Chatterton and Kay Francis, and I knew what they meant".^[56] She began to regret accepting the role; and, to add to her disappointment, she was not confident in the abilities of her leading man, James Davis in his first major screen role. She disagreed with amendments made to the script because of censorship restrictions and found that many of the aspects of the role that had initially appealed to her had been cut. The film was later described by Bosley Crowther as "interminable"; and he noted that "of all the miserable dilemmas in which Miss Davis has been involved ... this one is probably the worst". It failed at the box office, and the studio lost nearly one million dollars.^[57]

While making June Bride (1948), Davis clashed with co-star Robert Montgomery, later describing him as "a male Miriam Hopkins... an excellent actor, but addicted to scene-stealing".^[58] The film marked her first comedy in several years and earned her some positive reviews; but it was not particularly popular with audiences and returned only a small profit. Despite the lackluster box office receipts from her more recent films, in 1949, she negotiated a four-film contract with Warner Bros., which paid $10,285 per week and made her the highest-paid woman in the United States.^[59] Jack Warner refused to allow her script approval, however, and cast her in Beyond the Forest (1949). Davis reportedly loathed the script and begged Warner to recast the role, but he refused. After the film was completed, Warner released Davis from her contract, at her request. The reviews that followed were scathing; Dorothy Manners writing for the Los Angeles Examiner described the film as "an unfortunate finale to her brilliant career".^[60] Hedda Hopper wrote, "If Bette had deliberately set out to wreck her career, she could not have picked a more appropriate vehicle."^[61] The film contained the line, "What a dump!", which became closely associated with Davis after it was referenced in Edward Albee's Who's Afraid of Virginia Woolf?, and impersonators began to use it in their acts. In later years, Davis often used it as her opening line at speaking engagements.

Starting a freelance career (1949–1960)[link]

Davis posing as Margo Channing in a promotional image for All About Eve (1950). She is pictured with Gary Merrill, to whom she was married from 1950 to 1960 (her fourth and final husband)

By 1949, Davis and Sherry were estranged and Hollywood columnists were writing that Davis's career was at an end. She filmed The Story of a Divorce (released by RKO Radio Pictures in 1951 as Payment on Demand) but had received no other offers. Shortly before filming was completed, the producer Darryl F. Zanuck offered her the role of the aging theatrical actress Margo Channing in All About Eve (1950). Claudette Colbert, for whom the part had been written, had severely injured her back, and she was unable to continue. Davis read the script, described it as the best she had ever read, and accepted the role. Within days she joined the cast in San Francisco to begin filming. During production, she established what would become a lifelong friendship with her costar, Anne Baxter, and a romantic relationship with her leading man, Gary Merrill, which led to marriage. The film's director Joseph L. Mankiewicz later remarked, "Bette was letter perfect. She was syllable-perfect. The director's dream: the prepared actress."^[62]

Critics responded positively to Davis's performance and several of her lines became well-known, particularly, "Fasten your seatbelts, it's going to be a bumpy night." She was again nominated for an Academy Award and critics such as Gene Ringgold described her Margo as her "all-time best performance".^[63] Pauline Kael wrote that much of Mankiewicz's vision of "the theater" was "nonsense" but commended Davis, writing "[the film is] saved by one performance that is the real thing: Bette Davis is at her most instinctive and assured. Her actress–vain, scared, a woman who goes too far in her reactions and emotions–makes the whole thing come alive."^[64] Davis won a "Best Actress" award from the Cannes Film Festival, and the New York Film Critics Circle Award. She also received the San Francisco Film Critics Circle Award as "Best Actress", having been named by them as the "Worst Actress" of 1949 for Beyond the Forest. During this time she was invited to leave her handprints in the forecourt of Grauman's Chinese Theatre.

On July 3, 1950 Davis's divorce from William Sherry was finalized, and on July 28 she married Gary Merrill. With Sherry's consent, Merrill adopted B.D., Davis's daughter with Sherry, and in 1950, Davis and Merrill adopted a baby girl they named Margot. The family traveled to England, where Davis and Merrill starred in a murder-mystery film, Another Man's Poison (1951). When it received lukewarm reviews and failed at the box office, Hollywood columnists wrote that Davis's comeback had petered out, and an Academy Award nomination for The Star (1952) did not halt her decline.

Davis and Merrill adopted a baby boy, Michael, in 1952, and Davis appeared in a Broadway revue, Two's Company directed by Jules Dassin. She was uncomfortable working outside of her area of expertise; she had never been a musical performer and her limited theater experience had been more than 20 years earlier. She was also severely ill and was operated on for osteomyelitis of the jaw. Margot was diagnosed as severely brain damaged due to an injury sustained during or shortly after her birth, and was eventually placed in an institution. Davis and Merrill began arguing frequently, with B.D. later recalling episodes of alcohol abuse and domestic violence.^[65]

Few of Davis's films of the 1950s were successful and many of her performances were condemned by critics. The Hollywood Reporter wrote of mannerisms "that you'd expect to find in a nightclub impersonation of [Davis]", while the London critic, Richard Winninger, wrote, "Miss Davis, with more say than most stars as to what films she makes, seems to have lapsed into egoism. The criterion for her choice of film would appear to be that nothing must compete with the full display of each facet of the Davis art. Only bad films are good enough for her."^[66] Her films of this period included The Virgin Queen (1955), Storm Center (1956), and The Catered Affair (1956). As her career declined, her marriage continued to deteriorate until she filed for divorce in 1960. The following year, her mother died. During the same time, she tried television, appearing in three episodes of the popular NBC western Wagon Train as three different characters in 1959 and 1961.

Renewed success (1961–1970)[link]

Davis received her final Academy Award nomination for her role as demented Baby Jane Hudson in What Ever Happened to Baby Jane? (1962), opposite Joan Crawford.

In 1961, Davis opened in the Broadway production The Night of the Iguana to mostly mediocre reviews, and left the production after four months due to "chronic illness". She then joined Glenn Ford and Ann-Margret for the Frank Capra film A Pocketful of Miracles (1961) (a remake of Capra's 1933 film, Lady for a Day), based on a story by Damon Runyon. She accepted her next role, in the Grand Guignol horror film What Ever Happened to Baby Jane? (1962) after reading the script and believing it could appeal to the same audience that had recently made Alfred Hitchcock's Psycho (1960) a success. She negotiated a deal that would pay her 10 percent of the worldwide gross profits, in addition to her salary. The film became one of the year's biggest successes.^[67]

Davis and Joan Crawford played two aging sisters, former actresses forced by circumstance to share a decaying Hollywood mansion. The director, Robert Aldrich, explained that Davis and Crawford were each aware of how important the film was to their respective careers and commented, "It's proper to say that they really detested each other, but they behaved absolutely perfectly."^[68] After filming was completed, their public comments against each other allowed the tension to develop into a lifelong feud. When Davis was nominated for an Academy Award, Crawford contacted the other Best Actress nominees (who were unable to attend the ceremonies) and offered to accept the award on their behalf should they win. Davis also received her only BAFTA Award nomination for this performance. Daughter B.D. played a small role in the film and when she and Davis visited the Cannes Film Festival to promote it, she met Jeremy Hyman, an executive for Seven Arts Productions. After a short courtship, she married Hyman at the age of 16, with Davis's permission.

In September 1962, Davis placed an advertisement in Variety under the heading of "Situations wanted—women artists", which read, "Mother of three—10, 11 & 15—divorcee. American. Thirty years experience as an actress in Motion Pictures. Mobile still and more affable than rumor would have it. Wants steady employment in Hollywood. (Has had Broadway)."^[69] Davis said that she intended it as a joke, and she sustained her comeback over the course of several years. Dead Ringer (1964) was a crime drama in which she played twin sisters and Where Love Has Gone (1964) was a romantic drama based on a Harold Robbins novel. Davis played the mother of Susan Hayward but filming was hampered by heated arguments between Davis and Hayward.^[70] Hush... Hush, Sweet Charlotte (1964) was Robert Aldrich's follow-up to What Ever Happened to Baby Jane?, in which he planned to reunite Davis and Crawford, but when Crawford withdrew allegedly due to illness soon after filming began, she was replaced by Olivia de Havilland. The film was a considerable success and brought renewed attention to its veteran cast, which also included Joseph Cotten, Mary Astor and Agnes Moorehead. The following year, Davis was cast as the lead in an Aaron Spelling sitcom, The Decorator.^[71] A pilot episode was filmed, but was not shown, and the project was terminated. By the end of the decade, Davis had appeared in the British films The Nanny (1965), The Anniversary (1968), and Connecting Rooms (1970), but her career again stalled.

Late career (1971–1983)[link]

Davis and Elizabeth Taylor in late 1981 during a show celebrating Taylor's life

In the early 1970s, Davis was invited to appear in New York, in a stage presentation, Great Ladies of the American Cinema. Over five successive nights, a different female star discussed her career and answered questions from the audience; Myrna Loy, Rosalind Russell, Lana Turner, Sylvia Sidney, and Joan Crawford were the other participants. Davis was well received and was invited to tour Australia with the similarly themed, Bette Davis in Person and on Film, and its success allowed her to take the production to the United Kingdom.^[72]

In 1972, she played the lead role in two television films that were each intended as pilots for upcoming series for NBC, Madame Sin with Robert Wagner, and The Judge and Jake Wyler, with Joan Van Ark, but in each case, NBC decided against producing a series. She appeared in the stage production, Miss Moffat, a musical adaptation of her film The Corn is Green, but after the show was panned by the Philadelphia critics during its pre-Broadway run, she cited a back injury and abandoned the show, which closed immediately. She played supporting roles in Burnt Offerings (1976) and The Disappearance of Aimee (1976), but clashed with Karen Black and Faye Dunaway, the stars of the two respective productions, because she felt that neither extended her an appropriate degree of respect, and that their behavior on the film sets was unprofessional.^[73]

In 1977, Davis became the first woman to receive the American Film Institute's Lifetime Achievement Award. The televised event included comments from several of Davis's colleagues including William Wyler who joked that given the chance Davis would still like to refilm a scene from The Letter to which Davis nodded. Jane Fonda, Henry Fonda, Natalie Wood and Olivia de Havilland were among the actors who paid tribute, with de Havilland commenting that Davis "got the roles I always wanted".^[74] Following the telecast she found herself in demand again, often having to choose between several offers. She accepted roles in the television miniseries The Dark Secret of Harvest Home (1978) and the theatrical film Death on the Nile (1978), an Agatha Christie murder mystery. The bulk of her remaining work was for television. She won an Emmy Award for Strangers: The Story of a Mother and Daughter (1979) with Gena Rowlands, and was nominated for her performances in White Mama (1980) and Little Gloria... Happy at Last (1982). She also played supporting roles in two Disney films, Return from Witch Mountain (1978) and The Watcher in the Woods (1980).

Davis's name became well-known to a younger audience when Kim Carnes's song "Bette Davis Eyes" became a worldwide hit and the best-selling record of 1981 in the U.S., where it stayed at number one on the music charts for more than two months. Davis's grandson was impressed that she was the subject of a hit song and Davis considered it a compliment, writing to both Carnes and the songwriters, and accepting the gift of gold and platinum records from Carnes, and hanging them on her wall.^[75][76] She continued acting for television, appearing in Family Reunion (1981) opposite her grandson J. Ashley Hyman, A Piano for Mrs. Cimino (1982) and Right of Way (1983) with James Stewart. In 1983, she was awarded the Women in Film Crystal Award.^[77]

Illness, conflict and death (1983–1989)[link]

In 1983, after filming the pilot episode for the television series Hotel, Davis was diagnosed with breast cancer and underwent a mastectomy. Within two weeks of her surgery she suffered four strokes which caused paralysis in the left side of her face and in her left arm, and left her with slurred speech. She commenced a lengthy period of physical therapy and, aided by her personal assistant, Kathryn Sermak, gained partial recovery from the paralysis.

During this time, her relationship with her daughter, B. D. Hyman, deteriorated when Hyman became a born-again Christian and attempted to persuade Davis to follow suit. With her health stable, she traveled to England to film the Agatha Christie mystery Murder with Mirrors (1985). Upon her return, she learned that Hyman had published a memoir, My Mother's Keeper, in which she chronicled a difficult mother-daughter relationship and depicted scenes of Davis's overbearing and drunken behavior. Several of Davis's friends commented that Hyman's depictions of events were not accurate; one said, "so much of the book is out of context". Mike Wallace rebroadcast a 60 Minutes interview he had filmed with Hyman a few years earlier in which she commended Davis on her skills as a mother, and said that she had adopted many of Davis's principles in raising her own children. Critics of Hyman noted that Davis had financially supported the Hyman family for several years and had recently saved them from losing their house. Despite the acrimony of their divorce years earlier, Gary Merrill also defended Davis. Interviewed by CNN, Merrill said that Hyman was motivated by "cruelty and greed". Davis's adopted son, Michael Merrill, ended contact with Hyman and refused to speak to her again, as did Davis, who also disinherited her.^[78]

Davis in 1987, two years before her death

In her second memoir, This 'N That (1987), Davis wrote, "I am still recovering from the fact that a child of mine would write about me behind my back, to say nothing about the kind of book it is. I will never recover as completely from B.D.'s book as I have from the stroke. Both were shattering experiences." Her memoir concluded with a letter to her daughter, in which she addressed her several times as "Hyman", and described her actions as "a glaring lack of loyalty and thanks for the very privileged life I feel you have been given". She concluded with a reference to the title of Hyman's book, "If it refers to money, if my memory serves me right, I've been your keeper all these many years. I am continuing to do so, as my name has made your book about me a success."^[79]

Davis appeared in the television film As Summers Die (1986) and Lindsay Anderson's film The Whales of August (1987), in which she played the blind sister of Lillian Gish. The film earned good reviews, with one critic writing, "Bette crawls across the screen like a testy old hornet on a windowpane, snarling, staggering, twitching—a symphony of misfired synapses."^[80] Her last performance was the title role in Larry Cohen's Wicked Stepmother (1989). By this time her health was failing, and after disagreements with Cohen she walked off the set. The script was rewritten to place more emphasis on Barbara Carrera's character, and the reworked version was released after Davis's death.

After abandoning Wicked Stepmother and with no further film offers (though she was keen to play the centenarian in Craig Calman's "The Turn Of The Century" and worked with him on adapting the stage play to a feature length screenplay), Davis appeared on several talk shows and was interviewed by Johnny Carson, Joan Rivers, Larry King and David Letterman, discussing her career but refusing to discuss her daughter. Her appearances were popular; Lindsay Anderson observed that the public enjoyed seeing her behaving "so bitchy". He commented, "I always disliked that because she was encouraged to behave badly. And I'd always hear her described by that awful word, feisty."^[81]

Davis's tomb at Forest Lawn

During 1988 and 1989, Davis was feted for her career achievements, receiving the Kennedy Center Honor, the Legion of Honor from France, the Campione d'Italia from Italy and the Film Society of Lincoln Center Lifetime Achievement Award. She collapsed during the American Cinema Awards in 1989 and later discovered that her cancer had returned. She recovered sufficiently to travel to Spain where she was honored at the Donostia-San Sebastián International Film Festival, but during her visit her health rapidly deteriorated. Too weak to make the long journey back to the U.S., she traveled to France where she died on October 6, 1989, at 11:20 pm, at the American Hospital in Neuilly-sur-Seine. Davis was 81 years old. She was interred in Forest Lawn—Hollywood Hills Cemetery in Los Angeles, alongside her mother, Ruthie, and sister, Bobby, with her name in larger type size. On her tombstone is written: "She did it the hard way", an epitaph that she mentioned in her memoir Mother Goddam as having been suggested to her by Joseph L. Mankiewicz shortly after they had filmed All About Eve.^[82]

Reception and legacy[link]

In 1964, Jack Warner spoke of the "magic quality that transformed this sometimes bland and not beautiful little girl into a great artist",^[82] and in a 1988 interview, Davis remarked that, unlike many of her contemporaries, she had forged a career without the benefit of beauty.^[83] She admitted she was terrified during the making of her earliest films and that she became tough by necessity. "Until you're known in my profession as a monster, you are not a star", she said, "[but] I've never fought for anything in a treacherous way. I've never fought for anything but the good of the film."^[84] During the making of All About Eve, (1950) Joseph L. Mankiewicz told her of the perception in Hollywood that she was difficult, and she explained that when the audience saw her on screen, they did not consider that her appearance was the result of numerous people working behind the scenes. If she was presented as "a horse's ass ... forty feet wide, and thirty feet high", that is all the audience "would see or care about".^[85]

The Letter (1940)

While lauded for her achievements, Davis and her films were sometimes derided; Pauline Kael described Now, Voyager (1942) as a "shlock classic",^[86] and by the mid-1940s her sometimes mannered and histrionic performances had become the subject of caricature. Edwin Schallert for the Los Angeles Times praised Davis's performance in Mr. Skeffington (1944), while observing, "the mimics will have more fun than a box of monkeys imitating Miss Davis", and Dorothy Manners at the Los Angeles Examiner said of her performance in the poorly received Beyond the Forest (1949), "no night club caricaturist has ever turned in such a cruel imitation of the Davis mannerisms as Bette turns on herself in this one". Time magazine noted that Davis was compulsively watchable even while criticizing her acting technique, summarizing her performance in Dead Ringer (1964) with the observation, "her acting, as always, isn't really acting: it's shameless showing off. But just try to look away!"^[87]

She attracted a following in the gay subculture^[88] and was frequently imitated by female impersonators such as Tracey Lee and Charles Pierce.^[89] Attempting to explain her popularity with gay audiences, the journalist Jim Emerson wrote, "Was she just a camp figurehead because her brittle, melodramatic style of acting hadn't aged well? Or was it that she was 'Larger Than Life,' a tough broad who had survived? Probably some of both."^[83]

Her film choices were often unconventional; she sought roles as manipulators and killers in an era when actresses usually preferred to play sympathetic characters, and she excelled in them. She favored authenticity over glamour and was willing to change her own appearance if it suited the character. Claudette Colbert commented that Davis was the first actress to play roles older than herself, and therefore did not have to make the difficult transition to character parts as she aged.^[84]

Davis's signature and handprints at Grauman's Chinese Theatre

As she entered old age, Davis was acknowledged for her achievements. John Springer, who had arranged her speaking tours of the early 1970s, wrote that despite the accomplishments of many of her contemporaries, Davis was "the star of the thirties and into the forties", achieving notability for the variety of her characterizations and her ability to assert herself, even when her material was mediocre.^[90] Individual performances continued to receive praise; in 1987, Bill Collins analyzed The Letter (1940), and described her performance as "a brilliant, subtle achievement", and wrote, "Bette Davis makes Leslie Crosbie one of the most extraordinary females in movies."^[91] In a 2000 review for All About Eve, (1950) Roger Ebert noted, "Davis was a character, an icon with a grand style, so even her excesses are realistic."^[92] In 2006, Premiere magazine ranked her portrayal of Margo Channing in the film as fifth on their list of "100 Greatest Performances of All Time", commenting, "There is something deliciously audacious about her gleeful willingness to play such unattractive emotions as jealousy, bitterness, and neediness."^[93] While reviewing What Ever Happened to Baby Jane? (1962) in 2008, Ebert asserted that "no one who has seen the film will ever forget her."^[94]

A few months before her death in 1989, Davis was one of several actors featured on the cover of Life magazine. In a film retrospective that celebrated the films and stars of 1939, Life concluded that Davis was the most significant actress of her era, and highlighted Dark Victory (1939) as one of the most-important films of the year.^[95] Her death made front-page news throughout the world as the "close of yet another chapter of the Golden Age of Hollywood". Angela Lansbury summed up the feeling of those of the Hollywood community who attended her memorial service, commenting after a sample from Davis's films were screened, that they had witnessed "an extraordinary legacy of acting in the twentieth century by a real master of the craft", that should provide "encouragement and illustration to future generations of aspiring actors".^[96]

In 1977, Davis became the first woman to be honored with the AFI Life Achievement Award.^[97] In 1999, the American Film Institute published its list of the "AFI's 100 Years... 100 Stars", which was the result of a film-industry poll to determine the "50 Greatest American Screen Legends" in order to raise public awareness and appreciation of classic film. Of the 25 actresses listed, Davis was ranked at number two, behind Katharine Hepburn.^[98]

The United States Postal Service honored Davis with a commemorative postage stamp in 2008, marking the 100th anniversary of her birth.^[99] The stamp features an image of her in the role of Margo Channing in All About Eve (1950). The First Day of Issue celebration took place September 18, 2008, at Boston University, which houses an extensive Bette Davis archive. Featured speakers included her son Michael Merrill and Lauren Bacall.

In 1997, the executors of her estate, Michael Merrill, her son, and Kathryn Sermak, her former assistant, established "The Bette Davis Foundation" which awards college scholarships to promising actors and actresses.^[38]

Academy Awards milestones[link]

Davis in the trailer for Dark Victory (1939), in which she gave one of her ten Oscar-nominated performances

In 1962 Bette Davis became the first person to secure ten Academy Award nominations for acting. Since then only four people have equalled or surpassed this figure, Meryl Streep (with seventeen nominations and three wins), Katharine Hepburn (twelve nominations and four wins), Jack Nicholson (twelve nominations and three wins) and Laurence Olivier (ten nominations and one win).^[100]

Steven Spielberg purchased Davis's Oscars for Dangerous (1935) and Jezebel (1938) when they were offered for auction for $207,500 and $578,000, respectively, and returned them to the Academy of Motion Picture Arts and Sciences.^[101][102]

• 1934: Davis's performance in Of Human Bondage (1934) was widely acclaimed and when she was not nominated for an Academy Award, several influential people mounted a campaign to have her name included. The Academy relaxed its rules for that year only to allow for the consideration of any performer nominated in a write-in vote; therefore, any performance of the year was technically eligible for consideration. Given the well-publicized hoopla, some sources still consider this as a nomination for Davis; however, the Academy does not officially record this as a nomination.^[17]
• 1935: Won for Dangerous
• 1938: Won for Jezebel
• 1939: Nominated for Dark Victory
• 1940: Nominated for The Letter
• 1941: Nominated for The Little Foxes
• 1942: Nominated for Now, Voyager
• 1944: Nominated for Mr. Skeffington
• 1950: Nominated for All About Eve
• 1952: Nominated for The Star
• 1962: Nominated for What Ever Happened to Baby Jane?

Selected filmography[link]

• Of Human Bondage (1934)
• Dangerous (1935)
• The Petrified Forest (1936)
• Marked Woman (1937)
• Jezebel (1938)
• Dark Victory (1939)
• The Old Maid (1939)
• The Private Lives of Elizabeth and Essex (1939)
• All This, and Heaven Too (1940)
• The Letter (1940)
• The Little Foxes (1941)
• The Great Lie (1941)
• The Man Who Came to Dinner (1942)
• In This Our Life (1942)
• Now, Voyager (1942)
• Watch on the Rhine (1943)
• Mr. Skeffington (1944)
• All About Eve (1950)
• The Star (1952)
• Pocketful of Miracles (1961)
• What Ever Happened to Baby Jane? (1962)
• Hush... Hush, Sweet Charlotte (1964)
• The Nanny (1965)
• Death on the Nile (1978)
• The Whales of August (1987)

Notes[link]

References[link]

• Bret, David (2006). Joan Crawford: Hollywood Martyr. Carroll & Graf Publishers. ISBN 978-0-7867-1868-9. 
• Carr, Larry (1979). More Fabulous Faces: The Evolution and Metamorphosis of Bette Davis, Katharine Hepburn, Dolores del Rio, Carole Lombard and Myrna Loy. Doubleday and Company. ISBN 0-385-12819-3. 
• Chandler, Charlotte (2006). The Girl Who Walked Home Alone : Bette Davis, A Personal Biography. Simon and Schuster. ISBN 978-0-7432-6208-8. 
• Collins, Bill (1987). Bill Collins Presents "The Golden Years of Hollywood". The MacMillan Company of Australia. ISBN 0-333-45069-8. 
• Considine, Shaun (2000). Bette and Joan: The Divine Feud. Backinprint.com. ISBN 978-0-595-12027-7. 
• Davis, Bette; Michael Herskowitz (1987). This 'N That. G. P. Putnam's Sons. ISBN 0-345-34453-7. 
• Guiles, Fred Lawrence (1995). Joan Crawford, The Last Word. Conrad Goulden Books. ISBN 1-85793-268-4. 
• Haver, Ronald (1980). David O. Selznick's Hollywood. Bonanza Books. ISBN 0-517-47665-7. 
• Kael, Pauline (1982). 5001 Nights at the Movies. Zenith Books. ISBN 0-09-933550-6. 
• Ringgold, Gene (1966). The Films of Bette Davis. Cadillac Publishing Co. ISBN 0-8065-0953-8. 
• Shipman, David (1988). Movie Talk. St. Martin's Press. ISBN 0-312-03403-2. 
• Sikov, Ed (2007). Dark Victory: The Life of Bette Davis. Henry Holt and Company. ISBN 0-8050-7548-8. 
• Spada, James (1993). More Than a Woman. Little, Brown and Company. ISBN 0-316-90880-0. 
• Sperling, Cass Warner; Cork Milner and Jack Warner Jr. (1998). Hollywood Be Thy Name: The Warner Brothers Story. Prima Publishing. ISBN 0-8131-0958-2. 
• Springer, John; Jack Hamilton (1978). They Had Faces Then. Citadel Press. ISBN 0-8065-0657-1. 
• Staggs, Sam (2000). All About "All About Eve". St. Martin's Press. ISBN 0-312-27315-0. 
• Stine, Whitney; Bette Davis (1974). Mother Goddam: The Story of the Career of Bette Davis. W.H. Allen and Co. Plc.. ISBN 1-56980-157-6. 
• Wiley, Mason; Damien Bona (1987). Inside Oscar: The Unofficial History of the Academy Awards. Ballantine Books. ISBN 0-345-34453-7. 

Further reading[link]

• Davis, Bette (1962). The Lonely Life: An Autobiography. G. P. Putnam's Sons. 
• Stine, Whitney (1990). I'd Love to Kiss You: Conversations with Bette Davis. Pocket Books/Simon & Schuster. ISBN 0-671-74239-6. 

External links[link]

Biography portal

Wikiquote has a collection of quotations related to: Bette Davis

Wikimedia Commons has media related to: Bette Davis

• bettedavis.com, Operated by the Estate of Bette Davis
• Bette Davis at AllRovi
• Bette Davis at the Internet Broadway Database
• Bette Davis at the Internet Movie Database
• Bette Davis at the TCM Movie Database
• Bette Davis at TV Guide
• Bette Davis photographs and bibliography at virtual-history.com