Document DMwG9z2Mxe7xEDRorV8MO1LX5
pic*: me, -p and hoIJing fooJ in one free hand. I It- also wm'dle* fvirsx.-nd-, . uh ac nr bod* h.-ndt tmichin . overhanging nli).'Cts
hut not piri.idui't Mtvp.-usory suppott fur the body. lh:
someti
mio'cs over <!uxt m.laiiv.s with both forth,nl>*
raised and vttljlrc** hed above Ins lieail.
When
ii a Mel floor, he slides his knuckle walking
hands in fowl a.nl without lire: king mnlail with flic suit
As one foul swings for.va .Is, Die load is ls>rp.' almost ex
clusively nil the run'.r.il.iier.il foot and h.-iutch.
When moving rapidly on a dry substrate, lie raises into a
semi-erct quudruf'edaf position Me places his hands in
Ik walking pastures ulicuin the finger* are tightly Hexed ami
lha skin on the dorsum of umximul phalanges 11 -V makes con
tact with the substrate the hindlimb* are extended amt the ischial tiihcroiUics are well above the substrate. 1 he back and
head are ia. lined forwaids. The load is Write not only on the lateral asp-eels of the fret but also on the It.is as the subject
uses a "diagonal-sequent e, diagonal couplet* pail"'. He moves rapidly on a v. el lloor with alternate steps ol ihc
hindlimb* hut without raising hi* fist* limn the substrate.
Once he briefly slid his light hand forward* In a knuckle
walking posture while hi* lc.il hand remained a fist. The right hand bore only a moderate portion of the load because the
forelimbs remained protracted throughout the episode and
the body did not pass over the hands.
When leaning forwards to investigate or to pick up orally
objects on the floor and when descending head firsl from low
elevations, he consistently places his hands in fist walking
postures.
In summary, although Felix often places his hands in knuckle
walking po-.toies, he rarely support. a major portion of
his body weight on knuckle walking hands. Instead, he
place* hi* hand* in fist walking postures when support is
required for loads that arc significantly greater than the weight nf the foielind** themselves. By fi*t walking he probably
void* stressing metacaipoph.dHnge.il joints 11-V which, in
orangutans, lack special mnipliological adaptations for knuckle
walking of the kind possessed by gorillas nnd chimpanzees1 V
Before 1967, the primatologicnl and anthropological litera
ture was confusing nnd misleading about the terrestral hand
posture* ol great r>pc> because some authorities uncritically
nd sometimes erroneously designated orangutans, as well as
chimpanzees and gorillas, "knuckle walkers"'. One of us (ff .T.) recently described and classified* the variety
of hand |>osturcs often uscJ by captive orangutans during
terrestrial locomotion, and concluded that only chimpanzees
anvl gorillas should be designated knuckle walker*. He stated
that "orjn..ui;mi do not nnd, moreover, cannot assume the
digitigrmJc hand posture of the African ape*"''. This statement
should now be qualified and restricted to comparisons be
tween great .toe* engaged in semi-erect quadrupedal progres
sion with '11,11" and "diagonal sequence, duuoiial couplets"
gait?' on non l.ibricai'.'d substrates Although IVliv l as not
been ohs.'i ,vd to knuckle walk like an African ape or 10 plate his hands in knuckle walkm", p-osiurcl during "clutch walking"
as do chimpanzees and ;-, n'dl.iv, lie frequently tests his hands
on the middle segments of the fingers and Ihctcby Supports on llw'iu the vs v t.-hi of In, fore-limbs ,tnd perhaps also a port (tin
of bis lu.-d and upper luixO v\ light.
lie, .iiisv a lu :hly iidranet-d a: boreal climb,-i and arm swing, i
like an oi.incn.ut is able to place his li.ii'els in knuckle walkm';
po.lures sue csts that the an. eslois of the African apes might
have lv,:i simil.nly, ni !< a j-.rv.itcr estent, predisposed to
knutkk: v ilkioi', by (heir o.-.u s|Vcul a:boreal heritage.
M hi, in.i. olien r, sis Ii.s lie ml InU'ers in l.iviiftalivc kiuiekle
vs all in.- p ' i, s ;; v may [*, w itnessed in Cm i in pul .he speaker,
ami to- 'bill l.i.-incii Although th. lliuaih is usually uv d
pii*iiini' 11*> .i, a sii|i|ii'il`ii ; Mritl ilmm.; Iiimi.m l.inak'.
wallon-i (i .: mx. ilie I . i tli.il mail has a p..cdi-.po .ilio.i I'ei
- i I) pi., i i of li., .im.it di-si, 11 - V may at least :s piu-
v,', 'tie, I i 1 lution.ii v ini' i. in i > ai the facultative knuckle
v iMin.it ni i 'I all alt.iil
It may thu be argued that m m passed tht i.:.h t. >c nf arboreal tlimbing and suspensory pr.iuun; sou',*.** - ,t m,
advanced tftan the antibrnchi.itio-iists ,,n.l jnehr.--. "'.'u,.tt have admitted into their models' \ Am'il .i o.-dmii.d n,.*Jfl would evolve man, orangutans, or both Irom knuckl-- dkiux ancestor*. Such a theoretical possibility v.tmioi be on. J mi
on the ba*i of available evidence but v do rut thr>> c to subscribe to it.
Wc thank Wilh.mi Dnvisand Leland l .il'tamf ofih.-t t , -i,,,,
Zoologic.ll Park for their tuvUUiicO with vidcot. ,,n i ,.|,0
photography. This work wits supported by tin- f.
.,i
Science l utmdalion and by u IMIS research career doe; ,, i.c i award from the National Institute* of Health.
Department of Anthropology, Committee of Lrolutiortary Biology, and Biology Collegiate Dirhion, 112$ Eatt S9th Street, Unite!sity of Chicago, Chicago, Illinois 60637 .
kuistii 1 rn ii r
Chicago Zoological Park, DeookfieU, Illinois AU5I3. and Biology Collegiate Dirlsion, Unherslty of Chicago
Rfnmmin U fli e *
Received September 20, 1971.
' MiMcbrantl, M , Ainee. J. thys. imbn'i*'!.. 26 11*2 illff.-W 1 Tuttle, H. M., Amrr J Ph)S Anthropol. 20. 171 t, I -at, 7 > * 1 little, R. l.. Jufemr. 166. )) (19691.
* Tuttle, R. M., in The Chimpanzee (edit, by hourne, G H ' 2 (Kxrpcr. llnsle/New York. 19K)>.
* Straus, W. L.. Anie'r. J. Vhyt. Ambro/wl.. 17, 199 (19-tui
* Tuttkt, R. II.. Amer J. Phys. Anthropol.,lt, 171 (t,Jt>7j ' Turtle, R. II., Sri. J., 5A, No. J, 66 (1969).
i07
DSW 038085
IntcrpretvTtion of Persistence and Effects of Polychlorinated Biphenyls
in Birds
Peakxii and lancer' suggest thul polychlorinated bi.m-- iy!s
(RCIU) found in nature are derived only from highly
'.H
commercial mixiut.v such rn `Aroclor PSA' l(re,,|,.-s ...f
lower chlorinated mixtures have not been r.-pv u.d m \;"ie
of their predominant irvdiAtrial use. This may be <lu.` < Ub t
to utilization in envlosed systems or to more i.ipul iK-tab'ibv.'t
or ewtetion To demon.ti.uc which us,.!.! r,',im at .r
PCH feeding, pigeons (Cohonitin /nm) n.l / m. . ni
{Cotnrnix eotarni japonic o) have hcen t,-.l Art..I u' uvviu- s.
Twenly-fonr scpar.ilely caged feral ;
> ' 1 ad
libitum lot 2H da)s iulb wheat dressed with '>** t x- - .-
>ur
1242'. Sis control pigeons weic fed when At tin ; .i , : the
feedinR peiiv>d si\ lest birds were killo! to naMv.il ,b,' in.'ii
and the icin.utiJcr it-lurn-.d to u diet of mixed jy on :i"k in.e . x.
l>eforc b.'inkilled in batches of six at ?t V> a.nl I"'' <J.i,v
b.ightceii 4 week v>ld female Japanese ul weie , , .,1 in
tKilchv's of six. On; b.iich v as fed ntl liluium for .'l> d.,\$ wuti
`Artvlor I?d7` vhesxvd ftHHl at 2J(> ppm. i<ub :r "itb
'Atovloi 125V dtcswd food at 2512 P p 'll .Old the i, I uu,. c
batvh left as commix All hints werr kili.d 2 1 h .,l i ali-
drav ,,l vif dtessed l.iod. Ibis regno, a allows , ,n. j-.o , ,, "t
wills Other quail feeding cspci imvnts1. Ini.i.e 'i.il. b unr
death, breast nuisctc, hver. hiam and o uen' -I I -,i x . , |unf
rentes c,I fio,n pi 'ops Similar s.implcs '.etc n n> .,1 I mm
quad wiih ihc Mtbdilulion of heart for Im.ist umx.l,- A
STLCOPCB4021909
NAUJRt VOL 2V> MARCH 3 1972
TiLIf | kV.ft Total rca RasiMur-l (ppm.), found m Gua.l tissue
a|pl)iy.1ing V-.nH.tef 1342* and 'Atutoi 1
n 2!Or>(inv to# 20 (Jay*
1 i*sOC
liv*r Heart Brain Omental fai
Aroelo. 1-42" 17.0 *7.6* .1.2 i 0.7 7.8 + 1.7 123.1 17
'AriKltir 12W 78.1 1.9 7.0r l.l 7.7 4:2.3 30I3I
* AM Rgvrrt *ic mean of six bitdv + x.
portion of liver ws taken for preparation of mietosomc** and dclerminetion of cytochrome PJlt` and micrsstomal protein*. The remaining liver and tire other (issue* were uncled by the method of Taj lor rt at.* beTwo gas-liquid chromatographic anal) si* using a 3-foot 'Apiezon I.* column* Mire.
The complex nature of `Aroclor' mixtures makes the exprestin* of results difficult. Because gross chance* have taken ptac* in our experiment we have chosen the method of Risebrouch* from those puhlislted and compared the sum total of pttk height* with a l,l-di(4-chloroplicnyl)-2,2-dichloroethylce {DDE) standard, which in turn was related to a known weight f `Aroclor 1242'. The mean result* obtained from the batches of pigeons are shown in Fig. I and from quail in Tabic I. Regular checks made on excreta revealed no gross sanction of unchanged materi.il.
TIM presentation of changes in the gas-liquid chromaloptpbic pattern of PCB isomers is also difficult. Most workers have preferred to suggest that the patterns which they have obphtd resemble one of the commercial mixtures, leading to ilk end conclusion that PCBs remain unaltered in the bio sphere. Since we wished to determine whether hirds arc able U> metabolite PCBs we adopted the following method of ptmenUition for gas-liquid chromatographic data. A single Pk occurring in all chromatograms from fed birds was srlocied am) icpresented arbitrarily as UK) units. 1 he mean Of ptk height for each relative retention time from each batch of bird* was normalized to this peak. By this method an average residue ``spectrum" can be presented in the form of a
tig I M. in mini levels ul I't It pipeon (issues lollnmp
fit.Ill ;! <1 \lo.lnl I'-I.'1 ,|1 ;O.I n p III III llsl fill 7S Jjv*.
->&
) {* gfWl vv (Clult .,* .t( JltN'RVlJlrt
'mi
III!
<1 -L jno-
RltkluvLjl oft)fc\v<il iM
11*1 <4mi *>11 lag,
Juo-
i.--llLu,
lL
1 moitih alter "iihJrjw j| ofdressed din
i i
JtM
14* Mil u
tun
y jLjul "3
;
*uo
c b too
-
-U_
liiuneduuiv alter withdrawal i
of dressed dm
>
I ill 1 - i .. J
line reicifiKxi lime
Hs;. 2 Gas-liquid chromatographic "tpeclra", normalized to a common peak (vlippkd column) of `Aioclor 1241' standard and mean pigeon liver residue* following feeding at MX) p p m. in
diet Tor 28 day*.
tine diagram. Fig. 2 show* line diagrams from the lisrrs of
pigeons fed 'Aroclor 1242', ond Fig. 3 those from quail fed
'Aroclor 1754'. Other tissues gave similar results.
Total PCB residues immediately after withdrawal of treated
food show a similar distribution bclvvccn tissues to lliosc
in pigeons fed l,l-di(4-ch!orophcnyl)-2.2,2-tf ichlorocih.ine
(DDT)*. Allow ini; for the higher feeding level in the DDT
study initial PCB icsidues are live to ten times higher In tire
muscle and fat lltere is a steady logarithmic decay with half
lives. by inspection, of 50 days ami 123 days tv selectively. A her
all initial Luge drop in level* the liver icsnhtes also lotion ,x
logarithmic decay with a half-life of t-40 days 'these liyuies compare" with a statistically coin puled half-life of ?X d.iv*
fvir imr in all pu-eoii tissues (uneettainty in inletpieimp, t'L It
un.ilyltc.il results piecllides st.ile.iicul analysis), I'CTI lesuhie*
in the brain reach a maximum and then decay lop.iriihn u.ill)
with a half-life of fit days. I Ik- high User icmJuc level
immediately after wilhdiawal of food may he allrihm.ihie to
the ule at which liver icecivcv freshly absorbed PCD. and the
initial low brain level may icflctl the operation of the bloi-J-
biam barrier*.
total rc-iduov in quail fed 'Aii'clor 1242' (l..hk I) are
higher than those in pigeons in spite of a shorter fivdnn- period
at half llte level. Food coilsiimplivui per unit weight of quail,
however, is two to (luce fold In per than that - t pianos
Residues of 'Armior I2M* are gcncully twice tin<-e (>f
`Aroclor 17-12', suggesting that ilns mixture is more dulicult
to metabolize or excrete.
I he standard 1242 mixture gives a pattern dnnun.iied by a
few isomep. of low retention time on our g.is lupinl . hromatO-
pa.tph. 'I here is a eonsiileiable i h..o- e in pipes'll lot iv-idor
|).ltlellls (1 in. 3) with pc.ll.x of low Iv'lallVc' tellM'I'll times
1 -
a. I .............. .
I. .11, w- ,l(r , t|) 1.11 l a .1 I 4*
03W 038086
STLCOPCB4021910
M NATUflf V(H 7J* MAIICii J ^tTZ
hwwlMkl) .itko uitlhli.iujf cftfns <J (ltd
ToMo H M**n I W.xySi, Minos,.,* it r.ni . , i1 ,. ' ' '.Iimhi
Faip l*v*U am! l<,,v.<)a Lrv.ls (m /V|t-ixn Ixt) A.. 121? at bOO ppm tew Jli U. And Xilteil .i| 0 J;i. !*6 .md top li >* xT>.- ih*
' WnhO)) of Uu tKcO food
Day* to death
Control
0 24
34 164
Mean hsef MHitHomal ( >iiOuon;"
ws-iglrt
pTKll in 1***0 nnvul i; *
(| n4 t! ' live r liver
6.97 0. t! IV70* 1 07 2 40 0 78
10.75 J0 94 IV. 30 5 I.8U 15 47 t 1.70
g.7k * O.M 17 J! -t t M noo:l4t.
9.74 i 0 41 13.47 r 1.72 7 44 t. 0.43
6.50 r0.4 11.81 ) 1 40 6 73 i 0 81
ltd liter 1 .1 Juf* p | in
0
r. < . 1 8 1 l. 0 1
14 II t
i f> o :
* All figures arc mean of tie birds 11.
ToMo > Mean | ivuf Weiphta Mioroiouixl I'lolein Lovelt ( . , i..ama Tup Level* end Noxilue levele (or OukiI led Aror lor 1 74 e . .1 A.octr.,,
1 Jb4' at 7Ml ppm lor 70 Day* l>elo>* Ueoil,
01)1. Hclxihc mrttlHHi lime
Fig. J Gas-liquid chromatographic "spectra", normalized to a common peak (stippled column) of `Aroclor 1254' standard and mean quail liver residue* following feeding at 250 p.p.nt. for
20 day*.
('omiKnnxt fed
Mono liter Microsomal Cvrohrome , I *1 - I- v
wtip.hl
protein
I j,,i miH'l ' i. ,0..*-*
(g) mg g 1 liver r ' liver
cpm
Control
3.42 *0.22* 2J.0J* 2.27 14'*06l
`Aroclor 1242'4 52 10 37 31.62 2 I 27 22 14 2 83
`Aroclor 1234* 3.095 0.22 41.17 J 42 42.111 J-31
0 DO. '6 .`41179
treated food and virtually absent by the end of the experiment. These peaks seem to he readily removed from the body reveuling Ollier minor or less electron capturing constituents with a longer biological half-life. We cannot yei be certain that all these peaks arc chlorinated biphenyb present in the original mixture; some may be metabolites. Peaks with a higher reten tion time arc relatively enhanced over the experimental period. The patterns obtained after feeding quail `Aroclor 1242' have the same genernl features hut `Aroclor 1254' feeding produces kst aberration from the stundard.
The conclusion that the less chlorinated isomers are more readily metabolized is also supported by two analyses on a high resolution capillary column. A standard `Aroclor 1112' exhibited forty-four peaks and a considerable proportion could be assigned structures10. An extract of fat horn a pigeon killed immediately after withdrawal of dressed Torn! exhibited twenty nine (teaks. Most of the fifteen (teaks of lowest retention time (mainly dichloro and trichloro isomers) were absent from tin; extract. Peaks of longer retention time (mainly tarachloTO or higher chlorinated isomers) were present in the pigeon nnd many were relatively enhanced These results agree with pievioux work involving quail" and rats''
The hepatic mieioxomal Systran* responsible for riict.iholizinft foreign compounds are sensitise to induction by chlorinated hydrocarbons. The significance of induction Is nor yet understood hut it can bo used as an indicator of the biological activity of a compound. Increases in microsomal protein nnd cytochrome l\,,, arc central to microsomal induction and c have mcavurej these In indicate the interference by PCB residues in the normal physiology of the birds (Tables 2 and .1).
Pigeon h.er su'i -hl. remain elevated for at least two months
follow in;; withdrawal although hepatic microsomal protein returns to n. filial In contrast, t y t... hrome I'j>, l.vvl* are
still at ih; . lime the control level (> month* after ielmnin;;
to noim.J d.ct when Pill level* jrc a|*|>roxini iiety 2 ppm.
oihI compose d ir.'iuly nf lusher eh lot mated isomer *. s.nv.cslinji
(li.it evirer.ti lv lr level' ( < 1 p |> in.) of these isoin r* e iu*o
siyndie'.mt eU'Jiti*n of nti.rcivum.il cn/yinrs. Ouail led titlur
`Arovl.ir 1 2 2 or '1 'Sf show similar inerea-.es wlmli are- far
l.upcr ili.m lim e obtained from miiii1.ii cvpeiinvnrs urchin:: fllM ,md |i|)l . Am. lur l.'S-P priidueesevsnsieler.it). mvio
* Alt llgumere mean of *]* bird* i *..
induction than `1242', tupiKtrting tht suggestion above that tht more highly chlorinated biphenyls are the ohm potent inducer*
In spite of analytical ami interpretive diOVaili'.-v we can conclude that pigeons and quail metabolize pi .. . hi .ir mated blphcrtyb st a rate generally dependent on (he amount of
(chlorine in the molecule. Fmihec work in our laboratory
suggests that some lower chlorinated comer* are metabolized extremely rapidly. This could account for the almost tmiveral finding of residues of `Aroolur 1254 in spite of the majur production of '1242'. Bui biologicjl activity dep.nds on the highly chlorinated isomers, winch may need to be taken into account.
Wc thank Monsanto Chemicals l.td for `Arochu' miviurev and high resolution gas-liquid cluum.iiogi.iphy
Fist Infection < imtro! Laboratory, iWOtiit'} of dpi it n/ini r. Fisheries ami f:ui>tl,
lolnurlli, Surbiton, Slurry
S II VII i >
P. J. Ilrsvvs
OSw 038087
Received September 16; revised OelvHs i ?*, IWI
; Poalall. II .and I incer. J. I . tUouu,,,. 7 0 9'K(l')"'t)l
1 llunyan, P. J., lounwiul, M. (i .and Taylor, A (m ih,- pi<-*)
* Omura. I,, anil Sato, K.. J. Ih-il. C'h-m, 2.m. 2t7o (i-, n
A t.owiy. O It.. Itll.v l>-oueli, N. J,, 1 '.in /N J. Hull. < III m.. 1 v.t. 3r>3 043II.
t Taylor. A ,, Ke.i. It t . und hUrby. n It Simmons, 3. IP, and Tatum, J <3'(.<
! . . tncj K Mr i.t *i. < r
f-o/.i \ f H 9 W ll'JMi
Ch r^>l
M
(1V(.7|.
* Ri.chiuiii'.h. It. \V.. in
it! ft- ' *Nr (c.t it |) V II.. > <.* ( * . ,1 rK
Mill*-.. :si w.y. * 1 Cli .(lev ( . 1 ht"1 tu4 sl
'i. M. i) ItlIMt \
IWI)
It..,1**, s.. Ilunyan . P J . KrnntM'n, tt }> , ;i n \ t 1 A
7 a v f'.vf. .41'I'I !'`(..-Mi.itv/ .14, IUI 'J'l' ' t
Il.itkay. 1 .. Si h ull.i.
i|. 1||i| Vt.f ')
Si*w'ns. 1 > ., all.t U c fn, 1 > , J C hn**r f ' `1 f.d 1(- (f) f i t \ Koeman J II. I, N". scf IV IliJitiM M k . iml l ), \ r i l< II
A.ifwn-.IJI. II 'Ml'h.-Jl
" Gr.uu. I). I... Phillip.. W. I J ..i,l \ .U< muse. I> t l<,,U mp H. < .)lf )t; /nuiol , G, \1*' il > \ >
STLCOPCB4021911