Dustman 1985
Dustman 1985
R e c e i v e d 4 J a n u a r y 1985
T H E execution of complex behavtors undoubtedly depends jects LIberson [44] and Drechsler [27] noted that while SEPs
on a rather precise balance of excitatory and inhibitory ac- of young adults were localized to centro-panetal areas, SEPs
tivity within and between v a n o u s levels of the central nerv- of older mdtvlduals demonstrated a much wider spread and
ous system (CNS) It has been suggested that for elderly were found m occtpttal areas as well, findings that were at-
people, in comparison to young adults, the excitatzon/mMbi- tnbuted to decreased inhibitory function m old age [27]
tion balance has shifted such that the net electrical actw~ty of Stmdarly, Gaches [31] observed a wider dtstnbutton of alpha
their CNS tends to be relatively more excitatory or, con- actw~ty m the elderly The decreased localtzat~on of the
versely, relattvely less inhibitory Thus, for example, older brain's evoked and spontaneous electrical activity may re-
individuals are thought to have greater difficulty suppressing flect a breakdown of functional autonomy of cortical areas
or mhlbttmg extraneous st~muh Inlubitory deficits may con- resultmg from decreased central mMbmon
tribute to the reduced ability to attend and concentrate, the The present study is a continuation of our investigations
poorer memory, the decline m mental flexibility, and the of electrophysiolog~cal changes that accompany normal ag-
general slowing of perceptual performance that accompanies ing A battery of EEG and evoked potential measures were
aging [4, 11, 24, 30, 43] admmzstered to 40 young and 40 older individuals We report
Electrophyslologmal measures have provided presump- here the results of the EEG measures power spectral
tive evidence of central inhibitory deficmts in older people analysis which reflects abundance and amphtude of EEG
Components of sensory evoked cortical potentials have been within specified frequency bands, and cortical couphng
found to be larger for old than for young normal subjects [28, which provides an index of degree of correspondence of
29, 54] suggesting a relative inability of the CNS to dampen EEG waveforms from pairs of recording sites We hypoth-
or inhibit the effects of repeutlOUS sttmuh Others have ob- esized that if old age is associated with reduced mMbltory
served topographical spread of somatosensory evoked po- function and a 'breakdown of functional autonomy,' m con-
tentials (SEPs) and of alpha EEG m records of older sub- trast to the younger adults, (1) EEG power Ioadmgs of the
193
194 D U S T M A N ET AL
n.lO0 n
W ,o-q
,.-.q
,°q
,*o,
,°*o,
,.-°,
bo°,
~°-°,
,°-.,
~°. °1
~°o,
~°-.,
40 ¸ ~ m r/"
FIG 1 Mean IOglo EEG power for recordmgs from five scalp locations and for four frequency bands
Means were derived from power values of forty young (25-35 years) and forty older (55-70 years)
subjects
older individuals would be more evenly distributed (less var- an Age x Sex × Electrode × Band A N O V A Analyses were
ruble) across recordmg s~tes and (2) there would be a greater also done to determine ff age had a dLfferentlal effect on the
correspondence or congruence among E E G waveforms thus vanabdlty of power values across electrode sites A vana-
resulting m higher cortical couphng values The influence of bdlty score was determined for each subject by computing
gender on these measures was also mvesugated the standard deviation of the five power values, those from
Fz, Cz, C3, Pz, and Oz, for each of the bands The vanabdlty
METHOD scores were analyzed by an Age × Sex x Band A N O V A
Subjects t-Tests for independent means were used to compare vana-
bdlty scores of the 40 young subjects with those of the 40
F o u r groups of 20 normal volunteers participated in a older subjects for each of the four frequency bands
study of gender and age The groups comprised young
females and males aged 25-35 years and older females and
Corttcal Couphng
males aged 55-70 years All were screened with a health
quesUonna~re [1] for health problems that might adversely Cortical coupling was computed to measure phase rela-
influence their performance on electrophyslologlcal tests Uonshlps between E E G patterns from pairs of electrodes
All were judged to be m reasonably good health for their age, This time dependent measure is believed to reflect functional
all received a modest payment for thelr participation [42] communication, l e , information transmission between
brain areas [15, 65, 66]
EEG Recordmg Our procedures, modeled after those of Yagl et al [65],
used the Shannon-Weaver information transmission statistic
Subjects were seated in a comfortable, padded chlur in a
Txy [55] E E G from each electrode was dlgmzed at a 250/sec
darkened and electrically shielded room Disc electrodes
rate for 160 seconds Successive blocks of 400 digits were
were attached to the scalp with collodian at mldline (Fz, Cz,
used to calculate Txy The E E G at each sampling point for
Pz, and Oz) and left central (C3) sites, and referred to hnked
each of two inputs was classified as posture or negative with
earlobes (AI, A2) according to the International " 1 0 - 2 0 "
reference to a rather arbitrary baseline [62], and also as being
System [36] Subjects were requested to relax with eyes
larger or smaller than the previous dtgztai sample The sam-
closed whde three minutes of E E G was recorded on mag-
ple was then assigned to one of four categories positive and
neUc tape by a Hewlett Packard S-channel recorder The
nsmg, positive and falhng, negative and rising, negative and
E E G was amplified and traced on paper by a Grass (Model
falhng The correspondence between samples from two E E G
78B) 8-channel EEG/polygraph (band pass = 1-100 Hz)
signals was entered into a 4 x 4 contingency table Txy was
calculated from known probabd~t~es assocmted with the ta-
Power Spectral Analysts
ble's cell, row and column totals Cortical coupling values
Consecutive four second segments o f EEG, digitized at a can range from 0 00 (no relationship between E E G patterns)
128/sec rate, were displayed on a video terminal for visual to 2 00 (one E E G pattern can be predicted from the other)
lnspecUon and segments with obvious artifact were dis- An Age x Sex x Electrode pair A N O V A was computed on
carded E E G power was computed across a frequency band Txy values for the ten pa~rs of electrodes, an Age x Sex
of 3--15 Hz wzth 0 25 Hz resolution for each of the first 40 A N O V A was done for Txy values associated with each elec-
artifact-free segments using a fast Fourier approach [22,34] trode pair
and converted to ~tV2/(Hz/sec) units [61] A mean power
value for each 0 25 Hz point was obtamed by averaging RESULTS
across the 40 sets of values, ! e , those associated with the 40
four second segments Data reported in this paper are for Power Spectral Analysts
four frequency bands 5 25-7 00 Hz, 7 25-9 00 Hz, 9 25- Results from the Age x Sex × Area × Band A N O V A
11 00 Hz and 11.25-13 00 Hz (to be referred to as 5-7, 7-9, showed that gender did not have a slgmficant effect on E E G
9-11, and 11-13 H z bands) A mean was computed for the power nor did sex interact significantly with the other factors
eight 0 25 Hz power values comprising each band and con- ( p ' s > 0 30) However, E E G power differed slgmficantly
verted to log10 values [32] Power values were evaluated by across recording areas, F(4,312)=28 0. p < 0 0 0 1 Areas
PSA A N D C O R T I C A L C O U P L I N G O F E E G 195
TABLE 1
SIGNIFICANT RESULTS FROM AGE x AREA ANOVAs COMPUTED ON POWER
SPECTRAL DATA FOR FOUR FREQUENCY BANDS
ranked in terms of greatest to least power were Pz, Cz, Oz, 1 O0 [---'-']YOUNG
F z and C3 A post hot mean comparison test revealed that
each of the five means differed significantly from all other E " ~ OLD
area means Mean power was also reliably different for the 95
four frequency bands, F(3,234)=33 5, p < 0 001 Power was rr
largest for the 9-11 Hz band, next largest for 7-9 Hz and ,'.
smallest for 11-13 H z The means associated with the four 'i: 0 0 I
I
bands differed significantly from each other The Area × 8s
"
,°
Band interaction was also highly significant, F(12,936)=71 6, °.
-J ,°
p<0.001, demonstrating the E E G power for the four fre-
8O
quency bands was differentially distributed across recording
areas (see Fig l) In general, for mldhne recordings the fron-
tal and central sites generated more power within the two 75 PZ
lower frequency bands, 5-7 and 7-9 Hz, than did Pz and Oz Fz Cz Oz"
Pz and Oz, however, generated more power than F z and Cz
FIG 2 Mean log., EEG power for forty subjects aged 25--35 years
within the 9-11 and I 1-13 Hz bands Power associated with and forty aged 55-70 years The means mclude data for the 5-7.7-9,
C3 was larger for the two intermediate bands than for the 5-7 9--11 and 11-13 Hz bands
and 11-13 Hz bands
Age was not directly related to magnitude of E E G power,
F(1,78)=0 0 1 , p > 0 50 However, the Age x Area interaction
F(1,76)=33 7, p < 0 001 t-Tests (see Fig 3) demonstrated
was significant, F(4,312)=5 51, p < 0 001 Power values were
that the age-related difference in E E G power vanablhty oc-
larger for the young than for the older subjects at Cz and Pz
curred for each band There were no gender effects on power
while the opposite was true at F z , C3 and Oz locations (see
variability nor were there any significant interactions
Fig 2) The Age x Area x Band interaction was also sigmfi-
cant, F(12,912)=2 30, p < 0 01 The meaning of the interac-
Corttcal Couphng
tion was explored with Age x Area A N O V A s computed for
each frequency band E E G power o f the young and old An Age x Sex × Electrode Pair A N O V A , computed on
subjects was not different for any frequency band cortical couphng values, revealed significant maan effects for
( p ' s > 0 20), although there was a significant Age x Area in- Age, F(1,76)=442, p < 0 0 0 1 , and for Electrode Paar,
teraction for each band (Table l) Table 1 also shows that for F(9,684)=365 8, p<0.001, but not for Sex ( p > 0 20) F o r the
each frequency band there was a rehable difference among ten electrode pairs combined. E E G of the older subjects was
E E G power means for the five recording areas ( p ' s < 0 001) more highly coupled than that for the younger subjects With
Figure 3 portrays, for each frequency band, the distribution two exceptions, Cz-Pz vs Cz-C3 and C3-Oz vs Oz-Cz, the
of E E G power across electrode sites for the two groups of mean for each electrode pair was different from means for all
subjects other electrode pmrs Two significant mteractions were ob-
Variability of E E G power among the five electrode sites tained, Age × Electrode Pair, F(9,684)=, p<0.001, and Age
was computed for individual frequency bands for each sub- × Sex × Electrode Pair, F(9,684)=2 31, p < 0 02 To better
ject (see the Method section) The v a n a b d i t y measures were understand the interactions, an Age x Sex A N O V A was
evaluated with an Age x Sex x Band A N O V A A significant computed on coupling data for each electrode pmrmg Table
age effect was obtained with power being significantly less 2 shows that for all but two of the ten electrode pairs, cou-
variable among sites for the old than for the young subjects, phng values were reliably higher for the older than for the
1% DUSTMAN ET AL
- YOUNG TABLE 2
0. . . . . . ..O OLD MEANS, STANDARD DEVIATIONS, AND ANOVA F AND P VALUES
7-9 Hz FOR CORTICAL COUPLING DATA FROM 40 YOUNG (25-35YEARS)
5-7 Hz AND 40 OLDER (55-70 YEARS) ADULTS
younger SubJects @CO 001) There was no age dtfference for spread of EEG alpha from occlpltal cortex for old than for
Fz-C3 and Cz-C3 patnngs One margmally stgntficant Age x young subjects Similarly, SEPs of older mdlvlduals, m con-
Sex tnteractton was found, Cz-Pz couplmg values of young trast to those of younger SUbJeCtS, were more likely to be
males were shghtly larger than those of young females whde recorded at sites some distance from their pnmary source,
for the older group, means for males were smaller than those centro-panetal cortex [27,44] The increased SEP spread m
of females, F(1,76)=3 99, p=O 049 old age was attnbuted to reduced central mhlbltlon [27] We
In view of the significant age differences found for both concur with that interpretation and speculate that our results
EEG power vanablhty and cortical coupling values, the two also reflect an age-related shift m the bram’s exclta-
measures were correlated to determme their relatlonshlp tory/mhlbltory balance
For each SubJect a mean was computed from the four power There are neuroanatomlcal and neurochemlcal findings
vanability scores, I e , those for the four frequency bands, that are compatible with an hypothesis of reduced mhlbltlon
and a mean was computed from the cortical coupling values m elderly people Antenor cortical areas, believed to exert
for the ten pours of electrodes The correlation of these mean inhibitory control over the ascending reticular system [17,
values was -0 412 @<O 001,78&l Thus, reduced power var- 21,52,57] expenence substantial cell loss dunng old age [ 10,
lablllty across recording sites was associated with higher 13, 531 Old age IS also associated with reduced turnover of
cortical coupling values neurotransmitters [5,37], particularly the monoammes [5,46,
491 which include those whose predommant action 1s be-
lieved to be mhlbltory, I e , dopamme, norepmephnne and
DISCUSSION
serotonm [2, 18, 26, 45, 501
Two pnnclpal fmdmgs relating to adult aging emerged Sensory evoked potentials have been reported to reflect
from our study First, EEG power values were slgnlficantly mhlbltory deficits m old age Early occumng components of
less vanable across electrode sites for the 55-70 than for the visual (VEP) and somatosensory evoked potentials were
25-35 year old SUbJeCtS The effect was strong, being ob- larger for old than for young subjects (28,601 Late compo-
served for mdlvldual frequency bands as well as for bands nents of VEPs recorded from frontal-central areas were slg-
combined Second, coupling of spontaneous EEG was stgmf- mficantly larger for older than for younger mdlvlduals and
scantly higher for the old than for the young indicating an VEP amplitudes increased more rapldly for the elderly m
age-related mcrease m waveform congruity or slmllanty of response to increased flash intensity [29] The larger amph-
EEG from the several recordmg areas The two results may tudes and enhanced augmentation were believed to reflect a
reflect a common mechanism since across SubJects EEG decreased ability to mhlblt or dampen the effects of repeated
power vanability was slgmficantly related to cortical cou- sensory stimuli Down’s syndrome (DS) mdlvlduals who
plmg values (p<O 001). SUbJeCts whose power loadmgs were have an immature frontal cortex and bramstem [8] and re-
more homogeneous across recording areas tended to duced levels of serotomn [20] and possibly of dopamme [41]
demonstrate lugher cortical couplmg scores have been characterized as having central inhibitory deficits
Our fmdmgs of reduced vanablhty of EEG power and [25] They, hke normal older SubJects, have large evoked
higher corttcal couplmg values for the 55-70 year old sub- potentials [3, 9, 16, 58, 591 and VEPs that augment more
jects suggest there may be a breakdown of ‘fimctlonal au- strongly to bnghter flashes than those of age matched nor-
tonomy’ of areas wlthm the older bram such that it responds, mals [331 Additional evidence of a relationstip between
at least electncally, m a more homogeneous or global man- electrophyslology and mhlbltory function IS provided by
ner There have been other reports of increased congruity of studies of alcohohcs who show accentuated EP augmenta-
electrocortlcal activity among bram areas for older people tion [19,38, 391 Similar to elderly normals and DS mdwldu-
Gaches [31] observed that there was a greater antenor als, alcoholics have frontal cortical damage [23,43] and
PSA A N D C O R T I C A L C O U P L I N G O F E E G 197
analyses of their neurotransm~tter enzymes and metabohtes relaxed with eyes closed The effect of cogmUve tasks on
md~cate a reduced turnover of the monoanunes, particularly cortical couphng o f young and old subjects should provide
of serotonm and dopamlne [35,40] These findings appear to useful reformation regarding age-related changes m mntercor-
complement those of Begle~ter and coworkers who showed t~cal relationships
that chromc alcohol consumption by animals can result m a Our results offer vahdlty for cortical couphng procedures
hyperexc~table brmn [6,7] Couphng was slgmficantly higher for E E G recorded from
The effect of age on E E G power vanabdlty across record- adjacent electrodes than for E E G from more d~stant s~tes
mg s~tes was substantml, observed for each frequency band F o r example, the mean cortical couphng value for adjacent
w~th probabd~ty levels less than p =0 002 (F~g 3) These age pairs of mldlme electrodes (Fz-Cz, Cz-Pz, Pz-Oz) was 0 596,
differences were not the result of consistently h~gher or the mean for Fz-Pz and Cz-Oz was 0 254 and 0 124 for
lower E E G power loadmgs for the older group since the Fz-Oz Cortical areas that are close together m~ght be ex-
young and old subjects d~d not differ slgmficantly w~th re- pected to share more information than areas separated by
spect to mean E E G power for any recordmg area or fre- longer d~stances
quency band Despite the absence o f s~gmflcant age differ- The absence of gender differences for our elec-
ences m mean E E G power, there were suggestions of E E G trophys~ologlcal measures was somewhat surprising given
alpha slowing for the older individuals F o r the 9--11 Hz band that sex dtfferences have been reported for E E G charac-
the young subjects had shghtly larger power loadsngs at four teristics [14,48] and for evoked potential measures [56]
o f five recording sites whde the older subjects had larger However, E E G gender effects appear to occur p n m a n l y on
power loadmgs at all recording s~tes for the slower 7-9 Hz low voltage fast frequencies with females being more hkely
band (F~g 3) A slowing of E E G rhythms dunng adult agmg to have fast E E G records than males [14,48] Gender has not
has frequently been reported (e g , [47,63]) previously been investigated with cortical couphng tech-
The rehably h~gher cortical coupling values for the older tuques Our results suggest that commumcatlon between
subjects demonstrate a greater s~mdanty of their spontane- corhcai areas does not differ for males and females, at least
ous E E G among recordmg s~tes than was found for the for the brain at rest, ~ e , subjects relaxed, w~th eyes closed
younger adults The functional tmportance of th~s age- The posslbdlty that gender differences extst for a more active
dependent result ~s not known Based on their observations state of mental activity has been suggested by several E E G
that cortical couphng values can be altered by mampulatmg measures [51,64] Whether corhcal coupling techmques wdl
mode of cogmt~ve processing, Callaway and H a m s [15] hy- reveal sex differences as a function of task remains to be
pothesized that cortical couphng may reflect reformation investigated
transmission between brain areas While appropriate levels In summary, we report that two measures of E E G actlv-
of mterarea commumcat~on are undoubtedly essential for ef- ~ty, power spectral analys~s and cortical couphng demon-
ficient cognitive functioning, ~t may be that an excess of strate strong age effects Both indicate an increased s~mdar-
'cross talk' among areas interferes w a h these processes and ~ty of E E G activity among cortical areas m normal elderly
contributes to decreased cognmve function in elderly people people The results are compaUble wtth an hypothes~s of
[ 11,12] Our procedures devtated substantially from those of weakened inhibitory function in old age, perhaps the result
Cailaway and H a m s [15] as we d~d not ~mpose a cogmttve of neuroanatom~cal and neurochemlcal changes that are
load on our subjects other than having asked them to remam assocmted w~th growing old
REFERENCES
1 ASIM--Amertcan So~tet~ of Internal Medtcme Patients Per- i0 Bondareff, W The neural basis of aging In Handboo£ of the
sonal History Quesnonnatre Washington ASIM, 1978 Psychology of Aging, edited by J E Blrren and K W Schme
2 Austin. J H . E Connole, D Kett and J Collins Studies m New York Van Nostrand Reinhold, 1977, pp 157-176
aging of the brain V Reduced norepmephnne, dopamme, and 11 BotwmJck, J Aging and Behavior New York Springer, 1973
cychc AMP m rat brain with advancing age Age 1. 121-124, 12 Botwlmck, J Intellectual abdltles In Handboo£ of the Psy-
1978 chology of Aging. edited by J E Bwren and K W Schale New
3 Barnet, A B and A Lodge Chck evoked EEG responses in York Van Nostrand Reinhold, 1977, pp 580--605
normal and developmentally retarded infants Nature 214: 252- 13 Brody, H Aging of the vertebrate brain In Development and
255, 1967 Aging in the Nervous System, edited by M Rocksteln New
4 Bartus, R T and R L Dean Recent memory m aged non- York Academic Press, 1973, pp 121-133
human primates Hypersensitivity to visual interference dunng 14 Busse, E W and W D Obnst Pre-senescent electroencepha-
retention Exp Agmg Res 5: 385-400, 1979 Iographlc changes m normal subjects J Gerontol 20 315-320,
5 Beck, C H M Functional tmpllcatlons of changes m the senes- 1965
cent brmn A review Can J Neurol Set 5: 417--424, 1978 15 Callaway, E and P R Hams Couphng between cortical po-
6 Beglelter, H , V DeNoble and B Porjesz Protracted brain dys- tenuals from dd~ferent areas Sctence 183: 873-875, 1974
function after alcohol withdrawal m monkeys In B:ologwal 16 Callner, D A , R E Dustman, J A Madsen, T Schenkenberg
Effects of Alcohol, edited by H Beglelter New York Plenum and E C Beck Life span changes m the averaged evoked
Press, 1980, pp 231-249 responses of Down's syndrome and nonretarded persons Am J
7 Beglelter, H and B Projesz Persistence of brain hyperexclta- Ment Defic 82: 398-405, 1978
blhty following chronic alcohol exposure in rats In Alcohol 17 Campbell, B A , L D Lytle and G C Ftblger Ontogeny of
lntoxwat:on and Wlthdra~al, III, edited by M M Gross New adrenerg~c arousal and chohnerglc mhzbttory mechanisms m the
York Plenum Press, 1977, pp 209-222 rat Sczence 166. 635-637, 1969
8 Benda, C E Down's Syndrome New York Grune and Strat- 18 Carlson, N R Phys~ologyofBehawor Boston Allyn and Ba-
ton, 1969 con, 1977
9 Blgum, H B , R E Dustman and E C Beck Visual and 19 Coger, R W , A M Dymond, E A Serafetmldes, I Low-
somato-sensory evoked responses from mongoloid and normal enstam and D Pearson Alcohohsm Averaged visual evoked
chddren Electroencephalogr Chn Neurophystol 28: 576-585, response amphtude-mtenslty slope and symmetry m with-
1970 drawal Btol Pswhmtrv 11" 435-443, 1976
198 DUSTMAN ET AL
20 Coleman, M and D Mahanand Basehne serotomn levels In 45 McGeer, P L , J C Eccles and E G McGeer Molecular
Down's syndrome patients In Serotomn m Downs Syndrome, Neurobtologv of the Mammahan Brain New York Plenum,
edited by M Coleman New York Amencan Elsevier Publish- 1978
mg C o , 1973, pp 5-24 46 McGeer, P L and E G McGeer Chemistry of mood and
21 Como, P G , R Joseph, J D Flducla, J Siegel and J Lukas emotion Annu Rev Psychol 31" 273-307, 1980
Visually evoked potentials and after-discharge as a function of 47 Obrlst, W D Problems of aging In Handbook of Electroen-
arousal and frontal lesion m rats Soc Neurosct Abstr 5. 202, cephalography and Chmcal Neurophvstologx vol 6, Part A,
1979 edited by G E C h a t n a n a n d G C Lmry Amsterdam Elsevier,
22 Cooley, J W and J W Tukey An algonthym for the machine 1976, pp 275-292
computation of complex founer series Math Comput 19: 297- 48 Obnst, W D and E W Busse The electroencephalogram in
301, 1965 old age In Apphcattons of Electroencephalographx m Psvcht-
23 Courvllle, C B Effects of Alcohol on the Nervous System of ato' edited by W Wdson Durham Duke University Press.
Man Los Angeles San Lucas, 1955 1965, pp 185-205
24 Cralk, F I M Age differences m human memory In Hand- 49 Pradhan, S N and S Pradhan Development of central neuro-
boo~ o/the Psychology of Agmg, edited by J E Blrren and K transmitter systems and ontogeny of behavior In Btogent~
W Schale New York Van Nostrand Reinhold, 1977, pp 384-- Ammes In Development. edited by H Parvez and S Parvez
420 Amsterdam Elsevier/North Holland Biomedical Press, 1980,
25 Diamond, S . R S Balvm and F R Dmmond lnhibmon and pp 641-662
Chowe New York Harper and Row, 1963 50 Reader, T A The effects of dopamme, noradrenahne and
26 Douglas, W W Histamine and antihistamines, 5-hy- serotomn in the visual cortex of the cat Erpertentla 34 1586-
droxytryptamme and antagonists In The Pharmacologwal 1588, 1978
Basis of Therapeutws. edited by L S Goodman and A Gilman 51 Rebert, C S and R A Mahoney Functional cerebral asym-
New York Macmillan, 1975, pp 590-629 metry and performance III Reaction time as a function of task,
27 Drechsler, F Quantitative analysis of neurophyslologlcal proc- hand, sex, and EEG asymmetry Psvthophvsloh~gv 15. %16
esses of the aging CNS J Neural 218. 197-213, 1978 1978
28 Dustman, R E and E C Beck The effects of maturation and 52 Schelbel, M E and A B Schelbel The orgamzatlon of the
aging on the waveform of visually evoked potentials Elet- nucleus retlculans thalami A GolgJ study Brain Res 1 43--62,
troeneephalogr Chn Neurophystol 26. 2-11, 1969 1966
29 Dustman, R E and E W Snyder Life-span changes m visu- 53 Schelbel, M E and A B Scheibel Structural changes m the
ally evoked potentials at central scalp Neuroblol Aging 2 303- aging brain In Aging. vol !, edited by H Brody, D Harman
308, 1981 and J M Ordy New York Raven Press, 1975, pp 11-37
30 Frolkis, V V and V V Bezrukov Aging of the Central Nerv- 54 Shagass, C and M Schwartz Age, personal|ty and somatosen-
ous System New York Karger, 1979 sory evoked responses Science 148 135%1361, 1965
31 Gaches, J Etude statistique sur les traces "alpha largement 55 Shannon, C E and W Weaver The Mathematical Theor, of
developpe" en fonction de l'age Presse Med 68" 1620-1622, Commumcanon Urbana, IL Umverslty ofllhnols Press, 1949
1960 56 Shearer. D E , N B Cohn, R E Dustman and J A
32 Glanmtrapam, D Scanning mechanisms and the EEG Elec- LaMarche Electrophyslologlcal correlates of gender differ-
troeneephalogr Chn Neurophystol 30" 13%!46, 1971 ences A review Am J EEG Tethnol 24" 95--107, 1984
33 Ghddon, J B , J Busk and G C Galbraath Visual evoked 57 Skinner, J E and C D Ymghng Central gating mechanisms
responses as a function of light intensity m Down's syndrome that regulate event-related potentials and behavior A neurolog-
and nonretarded subjects Psvchophystologv 12: 416--422, 1975 ical model of attention In Progress m Chmtal Neurophvs-
34 Gold, G and C M Rader DtgitalProcessmg of Signals New tologv, e d l t e d b y J E Desmedt Basel Karger, 1977, pp 28-68
York McGraw-Hall, 1969 58 Straumams, J J , C Shagass and D A Overton Somatosen-
35 Gottfnes, C G , L von K n o m n g and C P e m s Neurophys- sory evoked responses m Down syndrome Arch Gen Psxcht-
lological measures related to levels of 5-hydroxymdoleacetlc att3' 29" 544-549, 1973
acid, homovamlhc acid and tryptophan m cerebrospmal fluid of 59 Stranmams, J J , C Shagass and D A Overton Auditory
psychiatric patients Neuropsychobtology 2: 1-8, 1976 evoked responses in young adults with Down's syndrome and
36 Jasper, H H The ten-twenty electrode system of the interna- idiopathic mental retardation Bud Psychiatry 6" 75-79. 1973
tional federation Electroencephalogr Chn Neuroph.~stol 10. 60 Straumams, J J , C Shagass and M Schwartz Visually evoked
371-375, 1958 cerebral response changes associated with chronic brain syn-
37 Kent, S Neurotransmitters may be a weak hnk m the agnng drome and aging J Gerontol 20" 498--506, 1965
brain's communication network Germtncs 31:105-111, 1976 61 Walter, D O On units and dimensions for reporting spectral
38 Knomng, L v o n Visual averaged evoked responses m patients intensities Electroencephalogr Chn Neurophvstol 24 486-487,
suffenng from alcoholism Neuropsychobtologv 2" 233-238, 1968
1976 62 Walter, D O Simplicity and difficulty of EEG analysis and a
39 Knomng, L yon and L Oreland Visual averaged evoked re- personal prognosis In EEG Informatws A Dtdacttt Revten of
sponses and platelet monamme oxldase activity as an md to Methods and Apphcanons of EEG Data Processing, edited by A
identify a risk group for alcoholic abuse A prehmmary study Remond New York Elsevier Scientific Publishing C o , 1977,
Prog Neuropsychopharmacol 2. 385-392, 1978 pp 407-413
40 Knomng, L von and C P e m s Biochem|stry of the 63 Wang, H S and E W Busse EEG of healthy old persons---a
augmenting-reducing response m visual evoked potentials longitudinal study I Dominant background activity and ocopl-
Neuropsychobiology 7" 1-8, 1981 tal rhythm J Gerontol 24. 419--426, 1969
41 Lake, C R , M G Zlegler, M Coleman and I J Kopm 64 Wogan, M , C D Kaplan, S F Moore and R Epro Sex differ-
Evaluation of the sympathetic nervous system m tnsomy-21 ences and task effects in laterahzatlon of EEG-alpha lnt J
(Down's syndrome) J Psychtatr Res 15" 1-6, 1979 Neurosct 8. 219-223, 1979
42 LaMarche, J A EEG and evoked potentml measures of age and 65 Yagl, A , L Bali and E Callaway Optimum parameters for the
sex differences in central nervous system processing Unpub- measurement of cortical coupling Phvstol Psvchol 4 33--38,
hshed doctoral dissertation, Utah State University, 1984 1976
43 Lezak, M D NeuropsvchologwalAssessment New York Ox- 66 Ymghng, C D Laterahzatlon of cortical coupling dunng com-
ford University Press, 1983 plex verbal and spatial behaviors In Language and Henn-
44 Llberson, W T Scalp dlstnbution of somato-sensory evoked sphertc Speclahzatton m Man Cerebral ERPs, edited by J E
potentials and aging Electromyogr Chn Neurophystol 16 221- Desmedt Basel Karger, 1977, pp 151-160
224, 1976