Abstract:

Almost all of the people experience a phantom limb after amputation. Sometimes so
real that they experience/sense(???????????) excruciating pain in the phantom. The
presence of phantom gives us the evidence of existence of plasticity in the adult
humans. Advanced technologies like fMRI, MEG help us track changes in the cortical
reorganization. Here, I talk about some of the experiences and experiments on
patients and the conclusions drawn about how the remapping in brain occurs.
Introduction:
The term phantom limbs was first given by Silas Weir Mitchell in 1871. During
wars, soldiers who underwent amputation often experienced phantom limbs. The
phantom limbs felt very real to the patients though the visual picture was
something different.
Earlier, it was thought that the much/most of the entire circuitry remains stable
throughout life under normal conditions (Wiesel and Hubel, 1963). Based on this, we
can say that no new neural connections are formed in the brain.
But, this turned out to be (not completely true)/false. Many experiments based on
the somatosensory map suggested that the neural connections do change and new
connections are formed even during old age (Wall,1977; Merzech et al.,1984; Wall,
1984; Pons et al.,1991) .
A Penfield map gives us the pictorial representation of the divisions in the motor
cortex and somatosensory cortex and tells us about the neural circuitry in the brain
of different body parts (Penfield and Rasmussen, 1950). The Penfield map is used
extensively to determine the remapping that has occurred in the brain after
amputation.

Body:
The neural circuitry that gives the feeling of pain, cold, hot,warmth is different from
that which gives the sensation of touch (Landgren, 1960; Kreisman and
Zimmerman, 1971).
Pons et al gave a map which tells us which part of the body is associated with what
part of the brain.

After deafferentation in monkey, it experiences the touch on both the face and the
amputated limb when stimulated on the face. This is due to change in the
somatosensory map after amputation (Pons et al.,1991).
To check whether the remapping also occur for pain, warmth, a stimulus test was
done by placing a drop of warm water on a patients face. They found that the
remapping also occurs in this case (Ramachandran et al., 1998).
The phantom limb vanishes over time. This could happen because there are
conflicting signals. The message from the motor cortex part of the brain is sent to
muscles in the hand regardless of the presence or absence of the limb
(Ramachandran and Rogers-Ramachandran, 1996). But a weird observation
regarding this is that in ~50% of the cases the limb becomes shorter and shorter
until only the palm and fingers remained and it remained attached and dangling
from the stump(reference). This phenomenon is called telescoping. This could
happen due to the over-expression of hand in the somatosensory cortex.
Simmel (1962) stated that children who/which are born with no limbs do not
experience phantom limbs. But this came into question when Weinstein et al. (1964)
studied some cases where children born without limbs were experiencing phantom
limbs which they were also able to move. Also another study of a 20-year old
woman who did not have limbs since birth suggested not only her experience of
phantom limb but also the ability to gesticulate those phantom limbs. But while
walking, her phantoms felt rigid, they werent moving (Ramachandran, 1993b). This
suggests that the origin of phantom did not come from her desire to be normal. This
could be because of the motor command sending signal to the phantom limb for
monitoring the reafference signal.

A poorly understood aspect of the phantom limb is the re-emergence of long lost
memories or the sensations that were felt just before amputation related to the
arm. They also felt clenching spasms in the hand and also digging in of nails. Once
the clenching stops, the digging in sensation also goes away. The reason is
unclear, nut a possible explanation is that when the motor cortex sends a signal to
clench the hand, there is an error feedback from proprioception and the signal is
normally damped. However, in the phantoms, such damping is not possible and
motor command is amplified even further, and this overflow of commands may be
experienced as pain.
The first clear demonstration of plasticity in the adult brain was given by Patrick
Wall and his co-workers(Wall, 1977). Output from single neuron in dorsal column
nuclei was taken. There was a big????? Change in the receptive field shortly after
partial denervation. They suggested that this change could be due to the activation
of the earlier silent synapses.
A study by Merzenich and his co-workers(1984) observed that if a monkey uses one
finger excessively for 90 min each day, within 3 months the cortex area pertaining
to that finger also expand but it does not go beyond ~1 mm. This 1-mm distance
was often thought to be fixed upper limit of the reorganization of sensory pathways
in adult primates. But an experiment by Pons et al (1991), suggested that it can go
beyond 1 mm. This was evident when they saw that touching the face stimulates
sensations in the arms.
Discussion and Conclusions:
Phantom limb studies gave an astounding proof of existence of plasticity in adult
primates. Extensive work has been done on phantom limbs. It gives a new fiend of
opportunity to investigate and understand how new connections emerge in the
brain, how the information from different sensory modalities interact and how the
brain changes its model in response to the external sensory inputs.
The cortical reorganization may serve as a compensatory mechanism by extending
zone of another senses to the one region of the amputated one. It is still debatable
whether phantom limb is a result of sprouting or unmasking.
Reference:
Kreisman, Norman R., and Irwin D. Zimmerman. "Cortical unit responses to
temperature stimulation of the skin." Brain research 25.1 (1971): 184-187.
Landgren, Sven. "Thalamic neurones responding to cooling of the cat's tongue."
Acta physiologica Scandinavica 48.2 (1960): 255-267.
Merzenich, Michael M., et al. "Somatosensory cortical map changes following digit
amputation in adult monkeys." Journal of comparative neurology 224.4 (1984): 591605.
Mitchell SW. Phantom limbs. Lippincott's Magazine 1871; 8: 563-69.

Penfield, Wilder, and Theodore Rasmussen. "The cerebral cortex of man; a clinical
study of localization of function." (1950).
Pons TP, Garraghty PE, Ommaya AK, Kaas JH, Taub E, Mishkin M. Massive cortical
reorganization after sensory deafferentation in adult macaques. Science.
1991;252:18571860.
Ramachandran, Vilayanur S. "Behavioral and magnetoencephalographic correlates
of plasticity in the adult human brain.[Review] " Proceedings of the National
Academy of Sciences of the United States of America 90 (1993b): 10413-20.
Ramachandran, Vilayanur S., and Diane Rogers-Ramachandran. "Synaesthesia in
phantom limbs induced with mirrors." Proceedings of the Royal Society of London.
Series B: Biological Sciences 263.1369 (1996): 377-386.
Ramachandran, Vilayanur S., and William Hirstein. "The perception of phantom
limbs. The DO Hebb lecture." Brain 121.9 (1998): 1603-1630.
Ramachandran, V. S., et al. "Illusions of body image: What they reveal about human
nature." The mind-brain continuum: Sensory processes (1996): 29-60.
Simmel, Marianne L. "The reality of phantom sensations." Social Research (1962):
337-356.
Wall, P. D., and J. H. Wolstencroft. "The Presence of Ineffective Synapses and the
Circumstances which Unmask Them [and Discussion]." Philosophical Transactions of
the Royal Society of London. B, Biological Sciences 278.961 (1977): 361-372.
Wall, Patrick D. "The painful consequences of peripheral injury." The Journal of Hand
Surgery: British & European Volume 9.1 (1984): 37-40.
Weinstein, Sidney, Eugene A. Sersen, and Robert J. Vetter. "Phantoms and somatic
sensation in cases of congenital aplasia." Cortex 1.3 (1964): 276-290.
Weiss, Samuel A., and Sidney Fishman. "Extended and telescoped phantom limbs in
unilateral amputees." The Journal of Abnormal and Social Psychology 66.5 (1963):
489.
Wiesel, Torsten N., and David H. Hubel. "Single-cell responses in striate cortex of
kittens deprived of vision in one eye." J Neurophysiol 26.6 (1963): 1003-1017.