The study of the neural basis of temporal
processing is in its infancy
However, it is arguably in the auditory domain
that timing is most prominent, owing to its importance in
vocalization and speech recognition.
Our brains measure time continuously. We
are aware of how long we have been doing a particular thing, how
long it has been since we last slept, and how long it will be until
lunch or dinner. We are ready, at any moment, to make complex
movements requiring muscle coordination with microsecond accuracy,
or to decode temporally complex auditory signals in the form of
speech or music. Our timing abilities are impressive, diverse
and worthy of investigation. But they are not very well
understood.
Many models of time perception have been put
forward (for example, see [1–3]), collectively
postulating a wide variety of different mechanisms.
Regardless of their diversity, the models all agree that temporal
information is processed in many ways: it is remembered, compared
to othertemporal information, combined with sensory information,
and used in the production of motor outputs.
The holy grail of timing research is to
understand the ‘time-dependent process’: a
mechanism equivalent to a piezoelectric crystal in a man- made
clock or the movement of a shadow on a sundial. This has proven
an elusive goal, to the extent that ideas about how this
mechanism might work remain near the level of conjecture.
Researchers have had great difficulty in pinning timing- related
activity in the brain to any specific type of function. This is
largely because mosttime measurement tasks draw upon more than
one process, making it difficult to tease the various components
apart.
In comparison with spatial stimuli, there is a
significant gap in our understanding of how the brain discriminates
simple temporal stimuli, such as estimating the duration of time
for which a light or tone is presented. Recent studies have begun
to examine the neural (Kilgard and
Merzenich, 2002; Hahnloser et al., 2002; Leon and Shadlen,
2003) and anatomical (Rao
et al., 2001; Lewis and Miall, 2003; Coull et al.,
2004) correlates of temporal processing. However,
the neural mechanisms that allow neural circuits to tell time and
encode temporal information are not clear. Indeed, it has
not yet been determined if timing across different time scales and
modalities relies on centralized or locally independent timing
circuits and mechanisms (Ivry and
Spencer, 2004).
Timing is critical in both the
discrimination of sensory stimuli (Shannon et al., 1995; Buonomano and Karmarkar,
2002; Ivry and Spencer, 2004; Buhusi and Meck, 2005)
and the generation of coordinated motor responses
(Mauk and Ruiz, 1992; Ivry, 1996;
Meegan et al., 2000; Medina et al., 2005). The
nervous system processes temporal information over a wide range,
from microsecondsto circadian rhythms (Carr, 1993; Mauk and Buonomano, 2004; Buhusi and
Meck, 2005).
Time and space are the fundamental dimensions
of our existence. Although space is gradually losing its value
in a world of computer networks, cellular phones and virtual
libraries, time is becoming the essence of our times, as is
reflected by ever increasing speed, rate of return and productivity
— concepts that are intrinsically related to time. Time is
also crucial for everyday activities, from our sleep–wake
cycle to walking, speaking, playing and appreciating music, and
playing sports. We can engage in these activities because, like
most animals, we process and use temporal information across a wide
range of intervals (FIG. 1) — in contrast to, for
example, the limited range of the light spectrum that we can
see.
Being able to tell the time is also advantageous
for gathering spatial information. Just as a position in space can
be triangulated by using distance to landmarks, the GLOBAL
POSITIONING SYSTEM (GPS) provides current position by triangulating
temporal information (the difference or coincidence in phase of
signals) from satellites. COINCIDENCE DETECTION is also used by
bats, owls and frogs to form an accurate, topographic
representation of space from INTERAURAL TIME DIFFERENCES1. For
these species, telling space is telling time. Timing and time
perception are fundamental to survival and goal reaching in humans
and other animals..
Many actions manifest precise timing. The
musicians in an orchestra time their movements to the gestures of
the conductor. The drag racer uses the countdown lights to
anticipate the start of a race. A pitcher must temporally
coordinate muscular activity across different joints to ensure that
the ball is delivered to a targeted region of the strike zone. One
basic question in motor control concerns how the timing of those
different types of actions is controlled and whether a common
process is invoked across different task domains so that such
temporal precision can be achieved (see Keele & Ivry,
1987).
Time perception is an ability that is
taken for granted, yet relatively little understood. Without it,
other cognitive functions, especially motor actions and visual
awareness, would be severely impaired. Basic tasks such as crossing
the road would be near impossible.