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Chapter 13 Spinal Control of Movement Answers to Chapter Review Questions Page 1 of 3 Question 1: What did Sherrington call the

e final common pathway, and why? nswer: Sherrington called the lower motor neurons of the spinal cord the final common pathway that controls !eha"ior# $hese motor neurons, also called the somatic motor neurons, directly command muscle contraction# $hey are the output of the motor system# %nputs to lower motor neurons include the sensory afferents entering the dorsal horn &pro"iding information a!out muscle length', the upper motor neurons in the motor corte(, and the interneurons within the spinal cord that participate in spinal motor programs# )egardless of the source of the input, the output is the lower motor neurons, the final common path# Question *: +efine, in one sentence, motor unit# ,ow does it differ from motor neuron pool? nswer: motor unit consists of one alpha motor neuron and all the muscle fi!ers that the alpha motor neuron inner"ates# $his is the elementary component of motor control# motor neuron pool consists of all the alpha motor neurons that inner"ate a single muscle# Question 3: Which is recruited first, a fast motor unit or a slow motor unit? Why? nswer: -ost muscles ha"e a range of motor unit $hese motor units are recruited in order of si.e/the smallest !eing recruited first and the largest last# $his e(plains why finer control is possi!le when muscles are under light loads than when they are under greater loads# Small motor units ha"e small alpha motor neurons and large motor units ha"e large alpha motor neurons# Small neurons are more easily e(cited !y signals descending from the !rain# Question 0: When and why does rigor mortis occur? nswer: $he stiffening of muscles after death is a condition 1nown as rigor mortis# -uscle contraction occurs !ecause of the interaction !etween myosin, the ma2or thic1 filament protein, and actin, the ma2or thin filament protein, during e(citation contraction coupling# $he heads of myosin filaments !ind to actin filaments and undergo a

conformational change# $his causes the thic1 filament to mo"e with respect to the thin filament, shortening the muscle fi!er during muscle contraction# $P is re3uired to release the myosin heads from the actin filament# When no $P is a"aila!le !ecause the tissue is dead, the attachment !etween the thic1 and thin filaments !ecomes permanent# Question 4: 5our doctor taps the tendon !eneath your 1neecap and your leg e(tends# What is the neural !asis of this refle(? What is it called? nswer: When your doctor taps the tendon !eneath your 1neecap, the tendon attached to the 3uadriceps muscle of your thigh is stretched# When this muscle is stretched, the muscle spindle afferents deli"er sensory feed!ac1 a!out the muscle length# $his causes the muscle to contract and your leg to e(tend# $his is a monosynaptic refle( arc in"ol"ing the spindle afferents that enter the dorsal horn and the motor neurons that control the muscle# $he 1nee6 2er1 refle( tests the intactness of the ner"es and muscles in this refle( arc# Question 7: What is the function of gamma motor neurons? nswer: lpha motor neurons inner"ate e(trafusal muscle fi!ers, causing muscle contraction# 8n the other hand, gamma motor neurons inner"ate the intrafusal muscle fi!er at the two ends of the muscle spindle# $he acti"ation of these fi!ers causes a contraction of the two poles of the muscle spindle, pulling the noncontractile e3uatorial region and 1eeping the %a a(ons acti"e# $he gamma motor acti"ity 1eeps the muscle spindle during muscle contraction under control# 8therwise, during muscle contraction, the muscle spindles would !ecome slac1 and insensiti"e to muscle length# Question 9: :enny, a character in Stein!ec1;s classic !oo1 Of Mice and Men, lo"ed ra!!its, !ut when he hugged them, they were crushed to death# Which type of propriocepti"e input might :enny ha"e !een lac1ing? nswer: :enny might ha"e !een lac1ing the propriocepti"e input of re"erse myotactic refle(# $he normal function of the refle( arc is to regulate muscle tension within an optimal range# %n e(treme circumstances, the refle( arc protects the muscle from !eing o"erloaded# $his type of propriocepti"e input is particularly important for the proper e(ecution of fine motor acts, such as the manipulation of fragile o!2ects with hands, which re3uires a steady, !ut not too powerful, grip# Question 1: :ist the components of the lateral and "entromedial descending spinal pathways# Which type of mo"ement does each path control?

nswer: $he components of the lateral descending spinal pathways are the corticospinal tract and the ru!rospinal tract# $he components of the "entromedial descending spinal pathways are the "esti!ulospinal tract, the tectospinal tract, the pontine reticulospinal tract, and the medullary reticulospinal tract# $he lateral pathways are in"ol"ed in the "oluntary mo"ement of the distal musculature# $he lateral pathways control the fine mo"ements of arms and fingers# $he "entromedial pathways control the posture of the head and nec1# Question *: 5ou are a neurologist presented with a patient who has the following symptom: an ina!ility to independently wiggle the toes on the left foot, !ut with all other mo"ements &wal1ing, independent finger mo"ement' apparently intact# 5ou suspect a lesion in the spinal cord# Where? nswer: :esions in the descending motor tracts, which originate in the upper motor system, can cause an a!normal <a!ins1i sign# $his was descri!ed !y =rench neurologist >oseph <a!ins1i in 1?@7# Scratching the sole of the foot from the heel toward the toes causes refle(i"e upward fle(ion of the !ig toe and an outward fanning of other toes# $he normal response to this stimulus for anyone older than * years is to curl the toes downward# Question 3: PA$ scans can !e used to measure !lood flow in the cere!ral corte(# What parts of the corte( show increased !lood flow when a su!2ect is as1ed to thin1 a!out mo"ing her right finger? nswer: When su!2ects are as1ed to mentally rehearse finger mo"ements without actually mo"ing the fingers, area 7 is acti"e !ut area 0 is not# $he portion of area 7 called S- sends a(ons that inner"ate distal motor units# $herefore, this area is li1ely to !e acti"e when rehearsing finger mo"ements rather than P- , which inner"ates pro(imal motor units# When su!2ects actually mo"e their fingers after rehearsing the mo"ement mentally, area 0 of the corte( registers increased !lood flow# $his is !ecause area 0 is in"ol"ed in e(ecuting mo"ements# Question 0: Why is dopa used to treat Par1inson;s disease? ,ow does it act to alle"iate the symptoms? nswer: $he organic !asis of Par1inson;s disease is a gradual degeneration of dopaminergic &+ ' neurons in the su!stantia nigra that pro2ect to the striatum# + normally facilitates the direct motor loop !y acti"ating cells in the putamen, which releases B:o from glo!us pallidus/induced inhi!ition# $he depletion of dopamine in Par1inson;s disease closes the funnel that feeds acti"ity to S- through the !asal ganglia and B:o# +opa is used to treat the depletion of dopaminergic input to the !asal ganglia caused !y Par1inson;s disease# +opa crosses the !lood6!rain !arrier and !oosts + synthesis in the dopaminergic neurons that remain in the su!stantia nigra# While this treatment alle"iates some of the symptoms, e"entually many

neurons are lost and dopa treatment is no longer effecti"e# +opa also has some trou!lesome side effects# Question 4: %ndi"idual <et. cells fire during a fairly !road range of mo"ement directions# ,ow might they wor1 together to command a precise mo"ement? nswer: Cpper motor neurons are located in cortical layer B of -1# :ayer B has a population of large pyramidal neurons called Betz cells. <et. cells were first descri!ed as a separate class of cells !y )ussian anatomist Bladimir <et. in 1?90# Single unit recordings in -1 !y Deorgopoulos and colleagues showed the following: i' -ost of the motor corte( is acti"e during e"ery mo"ement# ii' $he acti"ity of each cell represents a single "ote for a particular direction of mo"ement# iii' $he direction of mo"ement is determined !y a tally of the "otes registered !y each cell in the population# i"' lthough the population6coding scheme is hypothetical, e(periments on the superior colliculus conclude that a population code is used !y this structure to command precisely directed eye mo"ements# Question 7: S1etch the motor loop through the cere!ellum# What mo"ement disorders result from damage to the cere!ellum? nswer: (ons arising from layer B pyramidal cells in the sensorimotor corte(E frontal areas 0 and 7, somatosensory areas on the postcentral gyrus, and the posterior parietal areasEform a massi"e pro2ection to clusters of cells in the pons, the pontine nuclei, which in turn feed the cere!ellum# $he lateral cere!ellum pro2ects !ac1 to the motor corte( through a relay in the "entral lateral nucleus of the thalamus# $his completes the motor loop through the cere!ellum# +amage to the cere!ellum results in ata(ia, dysynergia, and dysmetria# ta(ia is a condition in which mo"ements !ecome uncoordinated and inaccurate# +ysynergia is characteri.ed !y the decomposition of synergistic multi2oint mo"ements# +ysmetria is characteri.ed !y clumsiness similar to that which accompanies ethanol into(ication# Question 1: ccording to the >ames6:ange and Fannon6<ard theories of emotion, what is the relationship !etween the an(iety you would feel after o"ersleeping for an e(am and your physical responses to the situation? nswer: ccording to the >ames6:ange theory of emotion, we e(perience emotion in response to physiological changes in our !ody, such as increased heart rate, inhi!ited digestion, and increased sweating# s a result

of the !odyGs response to the situation, the person !ecomes afraid# ccording to the Fannon6<ard theory, emotional e(perience is independent of emotional e(pression# $he threatening stimulus first causes a feeling of fear and the physiological reaction follows# Question *: ,ow has the definition of the lim!ic system and thoughts a!out its function changed since the time of <roca? nswer: =rench neurologist Paul <roca named the collection of cortical areas that form a ring around the !rain stem lim!ic lo!e &cingulated gyrus, medial temporal lo!e, including the hippocampus'# $here was no mention of emotionH the structures were primarily thought to !e in"ol"ed in olfaction# <y the 1@3Is, e"idence suggested that a num!er of lim!ic structures were in"ol"ed in emotion# merican neurologist >ames Pape. proposed an Jemotion systemJ on the medial wall of the !rain, which lin1ed the corte( with the hypothalamus &cingulated corte(, hippocampus, hypothalamus, anterior nuclei of the thalamus'# Pape. !elie"ed that damage to certain cortical areas caused profound changes in emotional e(pression with little change in perception or intelligence# Pape. proposed that acti"ity in the cingulated corte( adds emotional coloring# $he term lim!ic system was populari.ed in 1@4* !y merican physiologist Paul -ac:ean# ccording to -ac:ean, the e"olution of a lim!ic system ena!led animals to e(perience and e(press emotions# %t freed animals from the stereotypical !eha"ior dictated !y their !rain stem# Some of the components of the Pape. circuit are no longer thought to !e important for the e(pression of emotion, such as the hippocampus# %n addition, some structures in"ol"ed in emotion are also in"ol"ed in other functions, and some researchers 3uestion the utility of trying to define a single, discrete emotion system# Question 3: What procedures will produce an a!normal rage reaction in an e(perimental animal? ,ow do we 1now that the animals feel angry? nswer: $he remo"al of the cere!ral hemispheres produces a!normal rage reactions in e(perimental animals# A(periments performed in the 1@*Is showed a remar1a!le !eha"ioral transformation in cats or dogs when this procedure was performed# nimals that were not easy to pro"o1e prior to the surgery flew into a state of "iolent rage with the least pro"ocation after the surgery# Sham rage is o!ser"ed if the anterior hypothalamus is destroyed along with the corte(, !ut it is not seen if the lesion is e(tended to include the posterior half of the hypothalamus# $herefore, the posterior hypothalamus is particularly important for the e(pression of anger and aggression in animals and is normally inhi!ited !y the telencephalon# We

do not 1now whether or not the animals feel angry, !ecause feelings are su!2ecti"e e(periences that can !e reported "er!ally !y humans !ut not !y rats# Question 0: What changes in emotion were o!ser"ed following temporal lo!ectomy !y KlL"er and <ucy? 8f the numerous anatomical structures they remo"ed, which is thought to !e closely related to changes in temperament? nswer: Meuroscientists ,einrich KlL"er and Paul <ucy found that the !ilateral remo"al of temporal lo!es, also called temporal lobectomy, in rhesus mon1eys had a dramatic effect on the animalsG fear and aggression, which were decreased# $he animals were placid in the presence of humans and other animals that they normally fear, such as sna1es# $he animals showed a decrease in "ocali.ations and facial e(pressions typically associated with fear# %t appeared that !oth the normal e(perience and normal e(pression of fear and aggression were se"erely decreased# mygdala appears to !e a critical element in the !rain circuitry that processes fear and aggression# %t is thought to !e closely associated with changes in temperament# $he remo"al of amygdala reduces fear and aggression in e(perimental animals# Question 4: Why might performing !ilateral amygdalectomy on a dominant mon1ey in a colony result in that mon1eyGs !ecoming a su!ordinate? nswer: A"idence indicates that amygdala is in"ol"ed in aggressi"e !eha"ior# :esions of the amygdala may result in the flattening of emotion and other !eha"ioral a!normalities# <ilateral amygdalectomy in animals can profoundly reduce fear and aggression# $herefore, !ilateral amygdalectomy on a dominant mon1ey will ma1e the mon1ey placid and less difficult to challenge# s a result, the second mon1ey in the hierarchy will push the dominant mon1ey to a su!ordinate position# Alectrical stimulation of the amygdala may produce a state of agitation or affection aggression# Question 7: What assumptions a!out lim!ic structures underlie the surgical treatment of emotional disorders? nswer: $he assumption that the lim!ic system controls emotion led to the conclusion that people with emotional pro!lems can !e helped !y altering the system surgically# %n the 1@3Is, >ohn =ulton and Farlyle >aco!sen of the 5ale Cni"ersity reported that frontal lo!e

lesions had a calming effect in chimpan.ees# %t has !een suggested that frontal lesions ha"e this effect !ecause of the destruction of lim!ic structures, particularly in connection with frontal and cingulate corte(# &$his surgery is also associated with !lunted emotions, inappropriate !eha"ior, difficulty in planning and wor1ing toward goals, and difficulty in concentrating#' %n addition, reduced aggression in amygdalectomi.ed animals led some neurosurgeons to use this method in humans# Flinical reports claim considera!le success in reducing aggressi"e asocial !eha"ior, increasing the a!ility to concentrate, decreasing hyperacti"ity, and reducing sei.ures with this type of !rain surgery# Question 9: $he drug 1nown as is a serotonin6selecti"e reupta1e inhi!itor# ,ow does this drug affect a person;s le"el of an(iety and aggression? nswer: $he neurotransmitter serotonin is in"ol"ed in regulating aggressionH decreased serotonin is associated with an increase in aggression, and increased serotonin is associated with decreased aggression# $he lin1 !etween aggression and an(iety is not perfectly clear, !ut it is 1nown that serotonin antagonists increase aggressi"eness and agonists of the 46 ,$1< and 46,$1< serotonin receptors decrease an(iety and aggressi"eness in mice# %t is also 1nown from e(perimental wor1 in animals that an(iety and aggression increase and decrease together# is a selecti"e serotonin reupta1e inhi!itor &SS)%' that effecti"ely increases the amount of serotonin in the synaptic cleft !y pre"enting its reupta1e into the presynaptic element# $his increase in serotonin a"aila!ility is associated with decreased an(iety, so in addition to its use as an antidepressant, and other SS)%s are used as antian(iety agents# Question 1: Why do AADs with relati"ely fast fre3uencies tend to ha"e smaller amplitudes than AADs with slower fre3uencies? nswer: $he amplitude of the AAD signal depends on the synchroni.ation of the acti"ity of the underlying neurons# %f a group of cells are e(cited simultaneously, the tiny signals sum to generate one large surface signal# ,owe"er, when each cell recei"es the same amount of e(citation, !ut spread out in time, the summed signals are meager and irregular# %n this case, the number of acti"ated cells and the total amount of excitation has not changedH howe"er, the timing of the acti"ity has changed# $herefore, AADs with relati"ely fast fre3uencies tend to ha"e smaller amplitudes than AADs

with slower fre3uencies# Question *: $he human cere!ral corte( is "ery large and must !e folded e(tensi"ely to fit within the s1ull# What do the foldings of the cortical surface do to the !rain signals that are recorded !y an AAD electrode at the scalp? nswer: =or the most part, an AAD measures "oltages generated !y the currents that flow during synaptic e(citation of the dendrites of many pyramidal neurons in the cere!ral corte(# $he signal must penetrate se"eral layers of non6neural tissue, including the meninges, fluid, !ones of the s1ull, and s1in, to reach the electrodes# $he population of cells deep within the folds of the cortical surface contri!utes "ery little to the recorded AAD, which measures acti"ity only in the superficial layers of corte( close to the s1ull# Question 3: Sleep seems to !e a !eha"ior of e"ery species of mammal, !ird, and reptile# +oes this mean that sleep performs a function essential for the life of these higher "erte!rates? %f you do not thin1 so, what might !e an e(planation for the a!undance of sleep? nswer: Mo single theory of the function of sleep is widely accepted, !ut the most reasona!le ideas fall into two categories: theories of restoration and theories of adaptation# $he theory of restoration states that we sleep in order to rest and reco"er, and to prepare to !e awa1e again# $he theory of adaptation states that we sleep to 1eep out of trou!le, to hide from predators when we are most "ulnera!le or from other harmful features of the en"ironment, or to conser"e energy# A"en animals that ne"er rest, such as dolphins, gi"e each hemisphere a nap: a!out * hours asleep on one side, then 1 hour awa1e on !oth sides, * hours asleep on the other side, and so on for 1* hours e"ery night# Similarly, the !lind %ndus )i"er dolphin uses microsleeps of 067 seconds in duration, adding up to 9 hours in a *06hour day to rest its !rain# $his reinforces the importance of sleep !ut the reason for its importance remains un1nown# nd it is possi!le that sleep is simply a !yproduct of some other "ital process# Monethless, rats depri"ed of sleep lose weight in spite of increased food inta1e, !ecome wea1, accumulate stomach ulcers and internal hemorrhages, and in se"ere cases e"en die# $hey are una!le to regulate !ody temperature and meta!olic needs# Question 0: n AAD during )A- sleep is "ery similar to an AAD when awa1e# ,ow do the !rain and !ody in )A- sleep differ from the !rain and !ody when awa1e? nswer: )apid eye mo"ement sleep, or )A- sleep, is a state where the whole !ody &e(cept for the eye and respiratory muscles' is immo!ili.ed, and "i"id, detailed illusions called dreams are con2ured up# $he o(ygen consumption of the !rain is higher in )A- sleep than when the !rain is awa1e and concentrating on difficult mathematical pro!lems# Some areas, including primary "isual corte(, are e3ually acti"e in the two states# ,owe"er, e(trastriate cortical areas and portions of the lim!ic system are significantly more acti"e during )Asleep#

Fon"ersely, regions of the frontal lo!es are noticea!ly less acti"e during )A-# -ost of the !ody is incapa!le of mo"ing during )A- sleep, whereas the !ody can !e mo"ed normally when awa1e# $he paralysis that occurs during )A- sleep is almost a total loss of s1eletal muscle tone# $he muscles controlling eye mo"ement, the tiny muscles of the inner ear, and the muscles of respiration are the e(ceptions, as these are stri1ingly acti"e# +uring )A- sleep, the same core !rain systems that control the sleep processes of the fore!rain acti"ely inhi!it the spinal motor neurons, pre"enting the descending motor acti"ity from e(pressing itself as actual mo"ement# Question 4: What is a li1ely e(planation for the !rainGs relati"e insensiti"ity to sensory input during )A- sleep, compared to the wa1ing state? nswer: $he control of )A- sleep, as with the other functional !rain states, deri"es from diffuse modulatory systems in the core of the !rain stem, particularly the pons# $he diffuse modulatory systems control the rhythmic !eha"iors of the thalamus, which in turn controls many AAD rhythms of the cere!ral corte(H slow, sleep6related rhythms of the thalamus apparently !loc1 the flow of sensory information up into the corte(# Question 7: $he SFM recei"es direct input from the retina "ia the retinohypothalamic tract, and this is how light6dar1 cycles can entrain circadian rhythms# %f the retinal a(ons were somehow disrupted, what would !e the li1ely effect on a personGs circadian rhythms of sleeping and wa1ing? nswer: %nput from the retina to the suprachiasmatic nucleus &SFM' of the hypothalamus is essential and sufficient to entrain sleeping and wa1ing cycles to night and day# When retinal a(ons are disrupted, and this essential input to the SFM is a!sent, the sleep6 wa1e cycles cannot !e entrained !y light# Such an indi"idual would !e su!2ect to a free6 running cloc1, which would drift out of phase with the typical light or dar1 cycle !ecause a free6running cloc1 runs on a longer day than normal &*4 hours in the short6term, 3I637 hours in the long6 term'# Such an indi"idual would !ecome sleepy during the day and wa1eful at night, until the cycle drifted !ac1 into phase with the normal light or dar1 cycle# Question 9: What differences would there !e in the !eha"ioral conse3uences of a free6running circadian cloc1 "ersus no cloc1 at all? nswer: free6running circadian cloc1 still has certain alternating phases of sleep and wa1efulness, and other !eha"ioral and physiological cycles, such as !ody temperature, continue to alternate, although they may !ecome desynchroni.ed so that sleep6wa1e and !ody temperature cycle at their own pace, uncoupled# 8n the other hand, when the SFM is lesioned, circadian rhythms are a!olishedEthe periodicity is lost# =or e(ample, s3uirrels and mon1eys with no SFM ha"e persistent high6fre3uency rhythms of !oth !rain acti"ity and temperature with no e"idence of regular cycling#

Question 1: ,ow is it possi!le for a split6!rain person to spea1 intelligi!ly if the left hemisphere controls speech? %sn;t this inconsistent with the fact that the left hemisphere must direct motor corte( in !oth hemispheres to coordinate mo"ements of the mouth? nswer: Some midline features are represented in !oth sides of the !rain, such as the fo"ea, which is represented in !oth right and left hemispheres# -otor control of the mouth and laryn( may !e similarly represented on !oth sides of the !rain# %n addition, the motor system wor1s according to the population code rather than a strict one6to6one correspondence !etween neural acti"ity and neural output# $his may loosen the topographic relationships !etween motor corte( and motor output# =inally, the two hemispheres can communicate "ia the anterior commisure in split6!rain indi"iduals !ecause this su!cortical fi!er tract remains intact when the corpus callosum is se"ered# Question *: What can you conclude a!out the normal function of <roca;s area from the o!ser"ation that there are usually some comprehension deficits in <roca;s aphasia? -ust <roca;s area itself !e directly in"ol"ed in comprehension? nswer: =unctional locali.ation is an appealing and important concept that helps us understand how the ner"ous system processes sensory information and commands motor output# <ut it is important not to lose sight of the interconnectedness of "arious !rain structures# $he predominant function of <roca;s area is language e(pression, and this function is diminished when <roca;s area is lesioned# <ut the entire circuitry of language processing must also !e disrupted in the a!sence of this structure, affecting the functioning of the circuit as a whole, including Wernic1e;s area, which appears to mediate language comprehension# %n addition, <roca;s area does not ha"e easily discerna!le !oundaries, and stro1es rarely in"ol"e discrete regions of corte(# Such lesions may in"ol"e other !rain structures !eyond <roca;s area# =inally, it is 3uite possi!le that <roca;s area participates in language comprehension e"en though its main function is language e(pression# Question 3: Pigeons can !e trained to press one !utton when they want food and other !uttons when they see particular "isual stimuli# $his means the !ird can name things it sees# ,ow would you determine whether or not the pigeon is using a new languageE!utton6ese? nswer: ,uman language is creati"eEnew word com!inations and sentences are constantly !eing made, and the com!inations ha"e clear meaning according to the meaning of the indi"idual words plus the rules for arranging them# nimals use sym!ols to identify 1nown o!2ects, !ut creati"e use of sym!ols appears to !e limited# good way to determine whether an animal is capa!le of language is to see whether the animal can use the sym!ols it has learned in no"el com!inations, i.e#, com!inations that ha"e not !een

taught, to signify no"el stimuli# Question 0: What does the Wernic1e6Deschwind language6processing model e(plain? What data are inconsistent with this model? nswer: $he Wernic1e6Deschwind language processing model offers simple e(planations for 1ey elements of <roca;s and Wernic1e;s aphasias# lesion in <roca;s area seriously interferes with speech production !ecause the proper output signals for speech e(ecution can no longer !e sent to motor corte(# 8n the other hand, comprehension is relati"ely intact !ecause Wernic1e;s area is undistur!ed# lesion in Wernic1e;s area produces serious comprehension pro!lems !ecause this is the site where sounds are transformed into words# $he a!ility to spea1 is usually unaffected !ecause <roca;s area is a!le to dri"e the muscles re3uired for speech# $he following data is inconsistent with the Wernic1e6Deschwind model: i' Words do not ha"e to !e transformed into a pseudoauditory response in Wernic1e;s area# Bisual information can reach <roca;s area from "isual corte( without ma1ing a stop at the angular gyrus# ii' $he se"erity of <roca;s and Wernic1e;s aphasias depends on how much corte( is damaged !eyond the limits of <roca;s and Wernic1e;s areas# %n addition, aphasia is influenced !y damage to su!cortical structures, such as the thalamus and caudate nucleus, which are not in the model# When parts of corte( are surgically remo"ed, the resulting language deficits are usually milder than the deficits resulting from stro1e, which affects !oth cortical and su!cortical structures# iii' $here is significant reco"ery of language function after a stro1e# 8ther cortical areas apparently compensate for what is lost# i"' -ost aphasias in"ol"e !oth comprehension and speech deficits# Question 4: %n what ways is the left hemisphere usually language dominant? What does the right hemisphere contri!ute? nswer: ,emispheric dominance for language is !est illustrated !y the responses of people with a se"ered corpus callosum when "isual input is restricted to only one hemisphere# Mum!ers, words, and pictures "isually presented in the right "isual field &and thus the left hemisphere of the !rain' can !e repeated or descri!ed with no difficulty !ecause the left hemisphere is usually dominant for language# %n addition, o!2ects manipulated !y the right hand &!ut out of "iew of !oth eyes' can !e descri!ed# ,owe"er, such simple "er!al descriptions of sensory inputs are not possi!le for the right hemisphere# %f an image is shown only in the left "isual field or an o!2ect is felt only !y the left hand &and thus the right hemisphere', a split6!rain su!2ect is una!le to descri!e it# $his a!sence of response !y the right hemisphere is a conse3uence and demonstration

of the left hemisphere !eing language dominant# $he right hemisphere has language comprehension and can read and understand num!ers, letters, and short words as long as the re3uired response is non"er!al# Question 7: What e"idence is there that <roca;s area is not simply a premotor area for speech? nswer: Se"eral lines of e"idence indicate that <roca;s area is not simply a premotor area for speech# i' $he difference in the aphasic;s a!ility to use content words and function words suggests that <roca;s area and near!y corte( may !e particularly in"ol"ed in ma1ing grammatical sentences out of words# ii' Wernic1e suggested that the area damaged in <roca;s aphasia contains memories for the fine series of motor commands re3uired for articulating word sounds, and this theory is upheld !y some people# iii' Fomprehension is good with <roca;s aphasia !ut comprehension deficits can !e demonstrated !y tric1y 3uestions# i"' %n addition, patients sometimes ha"e considera!le anomia, suggesting that they ha"e pro!lems finding words as well as ma1ing the appropriate sounds#