REVIEW
A shot in the dark: the use of darkness to investigate visual development
and as a therapy for amblyopia
Clin Exp Optom 2013; 96: 363–372 DOI:10.1111/cxo.12084
Donald E Mitchell PhD MAppSc BSc LOSc
Department of Psychology and Neuroscience,
Dalhousie University, Halifax, Nova Scotia, Canada
E-mail: d.e.mitchell@dal.ca
Extended periods of complete darkness have long been used among other early experiential
manipulations to explore the role of visual experience in the development of the visual
pathways. In the last decade, short periods of darkness have been used to facilitate the
imposition of different or conflicting visual input each day to explore the manner by which
processes of perinatal development controlled by gene action are refined subsequently
by visual experience. Very recently, periods of complete darkness of intermediate length (10
days) have been shown to promote very fast recovery from amblyopia induced by prior
monocular deprivation (MD). When imposed immediately after a period of MD, in certain
circumstances, darkness appears to insulate against the development of amblyopia. It is
proposed that complete darkness may reverse maturation of many of the so-called braking
molecules in the visual cortex, so that it reverts to a more juvenile state.
Submitted: 12 March 2013
Revised: 16 April 2013
Accepted for publication: 26 April 2013
Key words: amblyopia, darkness, deprivation, stereopsis, visual acuity
Vision is the only sense for which its natural
physical input can be eliminated completely
for extended periods by immersion in a
light-free external environment, for which
rigorous safeguards are in place to avoid
accidental stray light. Although the external
environment for other senses can be made
stimulus-free, unintended sensory stimula-
tion can occur from internal sources. Con-
sider audition for example; even with a
noise-free environment, unavoidable audi-
tory stimulation can still arise through air-
borne sound or by bone conduction, as a
consequence of self-induced or regular auto-
nomic motor responses associated with res-
piration or cardiac activity.
The ability to eliminate light from the
environment permits study of how develop-
ment of the visual pathway proceeds after
birth in the complete absence of sensory-
(visually) driven neural activity. Arguably,
this is a similar situation to that prior to birth,
when development proceeds on the basis of
molecular cues that may be modulated by
neural activity of intrinsic origin, such as
intrinsic waves of activity in the retina
1,2
but
not on the basis of neural activity that is
visually driven. The initial developmental
papers of Hubel and Wiesel
3
in the 1960s
provided the inspiration for many studies
directed toward an understanding of the
role played by visual experience in postnatal
development. In addition to documentation
of the functional state of the neonatal
cortex, these studies included the many
newsworthy demonstrations of the conse-
quences of different forms of selected or
biased early visual exposure (where the
visual input represented an extreme of the
normal continuum as with input to just
one eye or with input restricted to a single
contour orientation, such as horizontal) that
were imposed exclusively (that is, provided
the only visual input) for extended periods
in early life.
4–7
As an aid to the interpretation
of studies of the effects of biased early visual
exposure, it was necessary to explore the
effects of complete visual deprivation. All
of these experiments were designed in the
context of contemporary views of the state of
the neonatal cortex and by the need to inves-
tigate potential roles that subsequent visual
experience could exert. As a consequence,
the periods of selective visual exposure (or
deprivation) were exclusive and episodes
of normal visual input were specifically
avoided. With the advent of new technolo-
gies, such as optical imaging of intrinsic
signals,
8,9
a consensus has begun to amass
on the development of the three major
response properties of cells in the primary
visual cortex (their specificity for ocular
dominance, orientation and direction of
motion) and the emergence of the separate
columnar maps for these properties onto the
cortex.
10
For ocular dominance and orienta-
tion selectivity, development appears to
occur in two phases, an initial experience-
independent stage that is completed at or
near birth, followed by a period of plasticity
during which neural circuits and cortical
maps are refined and subject to alteration by
visual experience;
10–13
however, at least in
ferrets,
14
directional selectivity may prove an
exception to the two-stage viewpoints as this
property emerges after birth without an
experience-independent stage.
The initial studies of visual cortical
development imposed periods of darkness
or exclusively abnormal visual input that
extended for many weeks or months;
however, the recognition of a substantial
anatomical and physiological scaffold at or
near birth suggests that subsequent visual
experience may serve more to refine a pre-
existing cortical architecture than to sculpt it
de novo. Examination of the former role for
experience invites different approaches that
include the use of mixed daily visual input,
in which normal and abnormal visual inputs
are pitted against each other on a daily basis.
This paper summarises the major findings of
studies from my laboratory of the effects of
mixed daily visual experience in early post-
natal life. Implementation of these studies
required the use of a secure darkroom facil-
CLINICAL AND EXPERIMENTAL
OPTOMETRY
© 2013 The Author Clinical and Experimental Optometry 96.4 July 2013
Clinical and Experimental Optometry © 2013 Optometrists Association Australia 363