Accomodations
Plan:
1. Accommodation
2. Land based animals and the shape changing lens
Accommodation is the process by which the vertebrate eye changes optical
power to maintain a clear image or focus on an object as its distance varies. In this,
distances vary for individuals from the far point—the maximum distance from the
eye for which a clear image of an object can be seen, to the near point—the
minimum distance for a clear image. Accommodation usually acts like a reflex,
including part of the accommodation-vergence reflex, but it can also be
consciously controlled. The main ways animals may change focus are:
-Changing the shape of the lens.
-Changing the position of the lens relative to the retina.
-Changing the axial length of the eyeball.
-Changing the shape of the cornea.
Focusing the light scattered by objects in a three dimensional environment into a
two dimensional collection of individual bright points of light requires the light to
be bent. To get a good image of these points of light on a defined area requires a
precise systematic bending of light called refraction. The real image formed from
millions of these points of light is what animals see using their retinas. Very even
systematic curvature of parts of the cornea and lens produces this systematic
bending of light onto the retina.
Due to the nature of optics the focused image on the retina is always inverted
relative to the object. Different animals live in different environments having
different refractive indexes involving water, air and often both. The eyes are
therefor required to bend light different amounts leading to different mechanisms
of focus being used in different environments. The air/cornea interface involves a
larger difference in refractive index than hydrated structures within the eye. As a
result, animals living in air have most of the bending of light achieved at the
air/cornea interface with the lens being involved in finer focus of the image.
Generally mammals, birds and reptiles living in air vary their eyes' optical power
by subtly and precisely changing the shape of the elastic lens using the ciliary
body.
The small difference in refractive index between water and the hydrated cornea
means fish and amphibians need to bend the light more using the internal structures
of the eye. Therefore, eyes evolved in water have a mechanism involving changing
the distance between a rigid rounder more refractive lens and the retina using less
uniform muscles rather than subtly changing the shape of the lens itself using
circularly arranged muscles.
[1]
Land based animals and the shape changing lens[edit]
Varying forms of direct experimental proof outlined in this article show that most
non-aquatic vertebrates achieve focus, at least in part, by changing the shapes of
their lenses.
What is less well understood is how the subtle, precise and very quick changes in
lens shape are made. Direct experimental proof of any lens model is necessarily
difficult as the vertebrate lens is transparent and only functions well in the living
animals. When considering vertebrates, aspects of all models may play varying
roles in lens focus. The models can be broadly divided into two camps. Those
models that stress the importance of external forces acting on a more passively
elastic lens and other models that include forces that may be generated by the lens
internally.
External forces[edit]
The model of a shape changing lens of humans was proposed by Young in a
lecture on the 27th Nov 1800.
[2]
Others such as Helmholtz and Huxley refined the
model in the mid-1800s explaining how the ciliary muscle contracts rounding the
lens to focus near
[3]
and this model was popularized by Helmholtz in 1909.
[4][5]
The
model may be summarized like this. Normally the lens is held under tension by
its suspending ligaments being pulled tight by the pressure of the eyeball. At short
focal distance the ciliary muscle contracts, stretching the ciliary body and relieving
some of the tension on the suspensory ligaments, allowing the lens to elastically
round up a bit, increasing refractive power. Changing focus to an object at a
greater distance requires a thinner less curved lens. This is achieved by relaxing
some of the sphincter like ciliary muscles allowing the ciliarly body to spring back,
pulling harder on the lens making it less curved and thinner, so increasing the focal
distance. There is a problem with the Helmholtz model in that despite
mathematical models being tried none has come close enough to working using
only the Helmholtz mechanisms.
Schachar has proposed a model for land based vertebrates that was not well
received.
[7]
The theory allows mathematical modeling to more accurately reflect
the way the lens focuses while also taking into account the complexities in the
suspensory ligaments and the presence of radial as well as circular muscles in the
ciliary body.
[8][9]
In this model the ligaments may pull to varying degrees on the
lens at the equator using the radial muscles, while the ligaments offset from the
equator to the front and back
[10]
are relaxed to varying degrees by contracting the
circular muscles.
[11]
These multiple actions
[12]
operating on the elastic lens allows it
to change lens shape at the front more subtly. Not only changing focus, but also
correcting for lens aberrations that might otherwise result from the changing shape
while better fitting mathematical modeling.
[6]
The "catenary" model of lens focus proposed by Coleman
[13]
demands less tension
on the ligaments suspending the lens. Rather than the lens as a whole being
stretched thinner for distance vision and allowed to relax for near focus,
contraction of the circular ciliary muscles results in the lens having less hydrostatic
pressure against its front. The lens front can then reform its shape between the
suspensory ligaments in a similar way to a slack chain hanging between two poles
might change it's curve when the poles are moved closer together. This model
requires precise fluid movement of the lens front only rather than trying to change
the shape of the lens as a whole.
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