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.