G rapevi ne d om esti cati on : w hy, how , w here and w hen?

D r Jos V oui l l am oz, U ni versi ty of N euchtel ,S w i tzerl and M ore detai l s about the present topi cwi l lbe gi ven i n the forthcom i ng book The G rape by Janci s R obi nson, Jul i a H ardi ng and Jos V oui l l am oz, S eptem ber 2012, P engui n P ress.
cate the w i l d grapevi ne? T he P al W hy dom esti eol i thi c H ypothesi s

Vi ti s vi ni fera subsp. si l vestri s) i sal i ana that natural l y cl i m bs up forest trees to The w i l d grapevi ne (
reach the canopy w here i t can produce fl ow ers that w i l ldevel op i nto berri es. These w el l -exposed berri es are very attracti ve to bi rds w ho propagate thei r seeds further, thus perpetuati ng the speci es. O ne can w onder w hy w oul d a P al eol i thi c m an or w om an brave the dangers of cl i m bi ng a tal lforest tree to reach those dark red berri es that are so attracti ve to bi rds but offer so l i ttl e food val ue? S erendi pi tous i nebri ati on m i ght be the answ er. Thi s has been cal l ed the P al eol i thi c H ypothesi s by eno-archaeol ogi st P atri ck M cG overn ( 2003) : attracted by the bri ghtl y col oured berri es or si m pl y mi mi cki ng bi rds, a group of earl y hum ans m i ght have pi cked som e bunches of grapes and w ere enti ced by the sugary, tart taste of the berri es. They w oul d then have gathered up severalcl usters i n an ani m alhi de, a w ooden contai ner or a rock crevi ce. A fter a few days, under the w ei ght of overl yi ng grapes, som e j ui ce w oul d have exuded from the berri es at the bottom . S i nce ferm entati on yeasts natural l y bl oom on the berri es ski ns and do exi st as spores i n the ai r, a l ow -al cohol i cwi ne coul d have been spontaneousl y produced. H avi ng eaten al lthe grapes, the P al eol i thi c m an or w om an w oul d have fortui tousl y tasted thi s beverage and a pl easant euphori a w oul d have hi m or her becom e obsessed by one si ngl e thought: to start agai n! W i thout proper contai ner, thi s S tone A ge B eauj ol ai s nouveau , as ni cknam ed by M cG overn, had to be dunk qui ckl y before i t turned to vi negar. Thi s w oul d change about 12000 to 10000 years ago w i th sedenti sm , the transi ti on from nom adi c to perm anent settl em ent that i si nti m atel y rel ated to the begi nni ngs of agri cul ture, a phenom enon know n as the N eol i thi c R evol uti on that al l ow ed the devel opm ent of dense popul ati ons and l ong-term food stori ng. These new l y settl ed popul ati ons needed to fi nd a w ay to secure grape suppl y.

H ow dom esti cati on w as m ade possi bl e? The H erm aphrodi ti c H ypothesi s I suggest to nam e the fi rst grape dom esti cati on event the H erm aphrodi ti c H ypothesi s , echoi ng the P al eol i thi c H ypothesi s of the di scovery of w i ne. B otani sts usual l y subdi vi de V i ti s vi ni fera i nto tw o subspeci es, V i ti s vi ni fera L. subsp. si l vestri s, the w i l d grape that natural l y occurs from P ortugalto Taj i ki stan as w el las al ong the m ai n conti nentalri vers of w estern E urope and i n northern A fri ca ( A rnol d et al . 1998) , and V i ti s vi ni fera L. subsp. vi ni fera ( al so cal l ed subsp. sati va) , the cul ti vated E urasi an grape.

N aturaldi stri buti on of the w i l d grape, V i ti s vi ni fera L. subsp. si l vestri s (C . C . G m el . ) H egi( from Zohary and H opf 2000) .
The m orphol ogi caltrai ts of the tw o subspeci es are often overl appi ng, and the val i di ty of thi s botani cal di sti ncti on coul d be questi oned because di fferences are m ost l i kel y the resul t of dom esti cati on by hum ans rather than evol uti onary di fferenti ati on ( Jacquat and M arti nol i1999; Zohary and H opf 2000; Thi s et al . 2006) . The m ost di scri mi nati ng character betw een w i l d and cul ti vated grapes i s fl ow er sex:

si l vestri si s di oeci ous, w hi ch m eans that al lthe fl ow ers on the sam e vi ne are ei ther stam i nate onl y( or
m al e) or pi sti l l ate onl y( or fem al e) , w hi l e vi ni fera i s herm aphrodi ti c, w i th fl ow ers havi ng stam ens and pi sti l . H ow ever, a ti ny percentage of si l vestri si s herm aphrodi ti c, w hi ch w as undoubtedl y the starti ng poi nt of grape dom esti cati on. In order to secure enough grape suppl y and avoi d the dangers of cl i m bi ng up trees, som e earl y M esol i thi c or N eol i thi c m an or w om an has taken i nto cul ti vati on w i l d grapevi nes ei ther by sow i ng seeds or by buryi ng cutti ngs i n the ground, w hat V i ti s vi ni fera spontaneousl y does i n nature ( thi si s cal l ed l ayeri ng) . If a m al e pl ant w as chosen, the vi ne w oul d never produce berri es and w oul d be qui ckl y abandoned. If a fem al e pl ant w hat chosen, the vi ne w oul d produce berri es onl yi f a m al e pl ant i si n the di rect nei ghbourhood for pol l i nati on, otherw i se the pl ant w oul d appear steri l e and w oul d be abandoned. If a herm aphrodi ti c pl ant w as chosen, grape harvest w oul d be secured each year ( berri es are usual l y produced by sel fi ng) and the vi ne w oul d be preserved. It i s therefore very l i kel y that grape dom esti cati on w as m ade from the ca. 2-3% of herm aphrodi ti cwi l d grapevi nes that are usual l y observed i n naturalpopul ati ons: thi si s our H erm aphrodi ti c H ypothesi s. The dom esti cated vi nes w ere then propagated by seed sow i ng or by cutti ngs, and those w i th l arger berri es, bi gger cl usters, hi gher sugar content or better arom as w ere acti vel y sel ected for m i l l enni a, w hi ch has l ead to the huge m orphol ogi caldi versi ty observed am ong the ca. 10000 grape vari eti es i n exi stence today.

W here w as the fi rst w i ne grape dom esti cated? The N oah H ypothesi s M ost am pel ographers ( gr. am pel os = grape, graphos = descri pti on) , archaeol ogi sts, botani sts and grape geneti ci sts agree that the ori gi ns of vi ni cul ture ( encom passi ng both vi ti cul ture and w i nem aki ng) are l ocated i n w hat I ni cknam e here the G rape s Ferti l e Tri angl e , a vast upl and regi on betw een the Taurus M ountai ns ( E astern Turkey) , the northern Zagros M ountai ns ( w estern Iran) and the C aucasus M ountai ns ( G eorgi a, A rm eni a and A zerbai j an) . P rom i nent R ussi an botani st and geneti ci st N i kol ai Ivanovi ch V avi l ov ( 1926) hypothesi zed that the area of greatest m orphol ogi caldi versi ty usual l y corresponds to the centre of ori gi n of a crop, w hi ch l ead hi s di sci pl e Al eksandr M i khai l ovi ch N egrul ( 1938) to the concl usi on that Transcaucasi a( the area betw een the B l ack and C aspi an seas extendi ng from the G reater C aucasus to the Turki sh and Irani an borders, and com pri si ng the R epubl i cs of G eorgi a, A rm eni a and A zerbai j an) or southern A natol i a( A si an part of Turkey) can be consi dered the

ti s hom el ands of the cul ti vated grape, si nce they are the areas show i ng the greatest di versi ty w i thi n Vi
scoveri ng w here and w hen the w i l d E urasi an grape w as fi rst vi ni fera. A recent D N A quest for di dom esti cated w as bapti sed w i th the m oni ker N oah H ypothesi s ( M cG overn 2003) , a paral l elw i th the E ve H ypothesi s traci ng back hum ani ty to a si ngl e ancestor through m i tochondri alD N A anal ysi s, as w el las a reference to N oah sl egend of pl anti ng the fi rst vi neyard on M ount A rarat ( a vol cani c dom e i n today s Turkey on w hi ch N oah s A rk i s sai d to have run aground but w here w i l d or cul ti vated grapes have never been observed) . A nal ysi s of rel atedness betw een subsp. si l vestri s and subsp. vi ni fera from al lE urasi a supported a geographi calori gi n of grape dom esti cati on i n the N ear E ast ( M yl es et al . 2011) and a focus on the G rape s Ferti l e Tri angl e show ed that the cl osest geneti c rel ati onshi p betw een l ocalw i l d grapevi nes and tradi ti onalcul ti vated grape vari eti es from southern A natol i a, A rm eni a and G eorgi a w as observed i n southern A natol i a, favouri ng the headw aters of Ti gri s and E uphrates R i vers i n the Taurus M ountai ns as the m ost l i kel y candi date for the l ocati on of the fi rst grape dom esti cati on event ( V oui l l am oz et al . 2004) .

The author Jos V oui l l am oz ( ri ght) together w i th P atri ck M cG overn (l eft) i n front of a w i l d grapevi ne on the shore of the headw aters of the Ti gri s Ri ver i n south eastern Turkey ( photo: A l iE rgl ) .
Thi si s consi stent w i th thi s area of form er M esopotam i a bei ng sti l lhom e to naturalpopul ati ons of w i l d grapevi nes, and w i th M cG overn s( 2003) reporti ng of abundant N eol i thi c( 10500-6000 before present, B P ) and E arl y B ronze A ge ( 10000-5000 B P ) evi dence for the presence of both w i l d and cul ti vated V i ti s

vi ni fera i n archaeol ogi calsi tes al ong the upper reaches of the Ti gri s and E uphrates R i vers, at the foot
of the Taurus m ountai ns ( ayn north-w est of D i yarbak r, H aci nebi , H assek H yk, K orucutepe, K urban H yk, Tepeci k, etc. ) . Thi s area al so i ncl udes the K aracada regi on i n the northern part of the Ferti l e C rescent that has been recentl y recogni sed by archaeol ogi caland geneti c studi es to be the core centre of dom esti cati on of severalfounder crops of agri cul ture: ei nkorn w heat, pea, chi ckpea, l enti l s, em m er and rye ( S al am i niet al . 2002) .

The Ferti l e C rescent ( dashed red l i ne) i s consi dered to be the cradl e of agri cul ture, w here founder cereal sl i ke ei nkorn w heat, em m er and rye and founder l egum es l i ke pea, chi ckpea and l enti l s w ere fi rst taken i nto cul ti vati on ( from S al am i niet al . 2002) . M aybe al so the w i l d grapevi ne?
The northern part of the Ferti l e C rescent w as al so show n by gl ottochronol ogy, the part of l i ngui sti cs deal i ng w i th the chronol ogi calrel ati onshi ps betw een l anguages, to be the pl ace of ori gi n of al lthe Indo-E uropean l anguages, w hi ch w as cal l ed the A natol i an Farm i ng H ypothesi s : usi ng unprecedented com putati onalm ethods deri ved from evol uti onary bi ol ogy to reconstruct the hypotheti calgeneal ogi cal tree of l anguages, G ray and A tki nson ( 2003) provi ded com pel l i ng cl ues that al lIndo-E uropean l anguages have stem m ed som e 8700 years ago from H i tti te, an exti nct l anguage of the south east A natol i a. Indo-E uropean l anguages w i l lthen have expanded from south east A natol i a som e 95008000 years ago al ong w i th the spread of agri cul ture, w hi ch speaks i n favour of the Ferti l e C rescent as the cradl e of m odern ci vi l i zati on. It seem s that al lcondi ti ons w ere m et i n south eastern A natol i a for successful l y engenderi ng grape dom esti cati on and earl ywi nem aki ng. In addi ti on, a cl oser l ook at the w ord for wi ne i n the m aj or Indo-E uropean l anguages that w ere i nvol ved i n earl y vi ni cul ture show s that they al lderi ve from the com m on P roto-Indo-E uropean ( the l anguage al l egedl y spoken by earl y popul ati ons of Transcaucasi a and eastern A natol i a) root *w oi -no or *w i n-o or *w i e-no. H ow ever, nonIndo-E uropean l anguages l i ke K artvel i an ( G eorgi an and M i ngrel i an) , S em i ti c( A kkadi an, U gari ti c and E arl y H ebrew ) and S em i to-H am i ti c( ol d E gypti an) al lpoi nt to an even m ore anci ent and unknow n com m on root, and no cl ear l i ngui sti c connecti on betw een these l anguage groups coul d be proposed. N everthel ess, som e G eorgi an schol ars have cl ai m ed that the ol dest root for the w ord wi ne i s the

K artvel i an vi no/

, sti l lused today i n m odern G eorgi an, w hi ch w oul d consti tute an i rrefutabl e

evi dence that G eorgi ai s the cradl e of vi ni cul ture ( G am krel i dze and Ivanonv 1990) .

The w ord for wi ne i n the m aj or l anguages i nvol ved i n vi ni cul ture. A l lIndo-E uropean w ords for w i ne deri ve from the com m on P roto-Indo-E uropean root *w oi -no or *w i n-o or *w i e-no. The asteri sk * i ndi cates a reconstructed form of an exti nct l anguage.
W hen w as the fi rst grapevi ne dom esti cated? W ewi l lprobabl y never know , but anal ysi s of archaeol ogi calgrape rem ai ns and j ars m i ght provi de som e cl ues. G rape rem ai ns found i n archaeol ogi caldi gs usual l y consi st of carboni sed pi ps and charred w ood, and thei ri denti fi cati on as w i l d( subsp. si l vestri s) or cul ti vated ( subsp. vi ni fera) i s probl em ati c. P i ps of subsp. si l vestri s are usual l y round w i th a short beak w hi l e pi ps of subsp. vi ni fera are pear-shaped w i th a w el l -devel oped beak ( S tum m er 1911; Terralet al . 2010) , but the carboni sati on process coupl ed w i th the enorm ous vari abi l i ty am ong both w i l d and cul ti vated groups m ake i t i m possi bl e to consi der pi ps m orphol ogy as a safe di agnosti c( Jacquat and M arti nol i1999; Zohary and

H opf 2000) . C arboni sed pi ps have been excavated from num erous prehi stori c archaeol ogi caldi gs i n E urope ( G reece, ex-Y ugosl avi a, Ital y, S w i tzerl and, G erm any, France and S pai n) ( Ri vera N unez and W al ker 1989) and i n south-w est A si a( Zohary and H opf 2000) , but these anci ent rem ai ns m ost l i kel y cam e from w i l d grape berri es that used to be col l ected l ong before dom esti cati on. A ccordi ng to G eorgi an am pel ographer R evaz R am i shvi l i , si x 8000 years ol d pi ps found i n the N eol i thi c si te i n S hul averi s-G ora i n the hi l l s south of Tbi l i si , one of the earl i est know n perm anent settl em ents i n G eorgi a, have a cul ti vated shape and al l egedl y consti tute evi dence of the fi rst dom esti cated grape

ti s vi ni fera subsp. vi ni fera ( M cG overn 2003) , but i denti fi cati on of these carboni sed rem ai ns seeds of V i
i s sti l lquesti onabl e. For archaeobotani sts D ani elZohary and M ari a H opf ( 2000) , grape pi ps found i n the C hal col i thi c( or C opper A ge, ca. 5700-5200 B P ) si te i n Tel lesh-S huna ( northern Jordan) consti tute the earl i est convi nci ng evi dence of grape cul ti vati on because w i l d Vi ti s vi ni fera subsp. si l vestri si s absent from Jordan today. H ow ever, even i fi ti s rather unl i kel y that these xeri c regi ons w ere hosti ng wi l d grapes som e fi ve or si xm i l l enni a ago, one coul d argue that they have now si m pl y di sappeared from thi s area. Less arguabl e are the evi dence found i n E arl y B ronze A ge ( ca. 5400-5200 B P ) si tes i n Jeri cho ( P al esti ne) , Lachi sh ( Israel ) , N um ei ra ( D ead S ea, possi bl y G om orrah?) , A rad ( Israel ) and K urban H yk ( near U rfa, south Turkey) , because not onl y carboni sed pi ps but al so charred grapevi ne w ood as w el las w hol e parched grape berri es w ere recovered, w hi ch provi des sol i d proof of cul ti vati on ( Zohary and H opf 2000) . H ow ever, earl y grapevi ne cul ti vati on m ust be m uch ol der than thi s archaeol ogi calevi dence, because the earl i est chem i calevi dence of w i ne m aki ng dates back to the 6th mi l l enni um B C . U si ng i nfrared spectrom etry to track dow n the presence of tartari c aci di nj ar deposi ts, w hi ch i s the si gnature of the presence of grapes, eno-archaeol ogi sts P atri ck M cG overn et al .( 1996) have found that w i ne w as al ready produced i n about 7400-7000 B P i n H aj j iFi ruz Tepe ( northern Iran, Zagros M ountai ns) ,l ocated i n the peri phery of m odern w i l d vi nes di stri buti on. These j ars cam e from a N eol i thi c resi dence: they w ere l ai d on thei r si de, em bedded i n the earthen fl oor of the ki tchen , and had cl ay stoppers: thi s m eans that they contai ned w i ne, not pl ai n grape j ui ce. Terebi nth tree resi n w as added to thi s retsi na-l i ke w i ne as a preservati ve. O ther evi dence of w i ne m aki ng has been found and thorough chem i cali nvesti gati on sti l lhas to be carri ed out, i n parti cul ar w i th sam pl es from S hul averi sG ora i n G eorgi a. V ery recentl y, excavati ons i n a cave at A renii n southeastern A rm eni a reveal ed the ol dest w i ne m aki ng i nstal l ati on, confi rm ed by l i qui d chrom atography and m ass spectrom etry anal yses, and dati ng from the C hal col i thi c peri od, som e 6000 B P ( B arnard et al . 2011) .

W i ne j ar found i n a N eol i thi c ki tchen i n H aj j iFi ruz Tepe ( northern Iran, Zagros M ountai ns) , dated 7400-7000 B P ( after M cG overn et al . 1996)
C oncl usi on Al though geneti cs, archaeol ogy and l i ngui sti cs tend to poi nt to southern A natol i a as the possi bl e cradl e of grape dom esti cati on, Transcaucasi a sti l lrem ai ns a seri ous candi date: anci ent grape rem ai ns w ere excavated from N eol i thi c archaeol ogi calsi tes i n C hokh i n D agestan, i n S hom u-Tepe i n A zerbai j an as w el las i n S hul averi s-G ora ( near Tbi l i si )i n G eorgi a( M cG overn 2003) , and chem i cal evi dence w as recentl y provi ded for w i ne producti on i n A reniw i ne regi on ( south east of Y erevan) i n A rm eni a around 6000 B P ( B arnard et al . 2011) .

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understand seed dom esti cati on syndrom e and revealori gi ns of anci ent E uropean cul ti vars' , A nnal s of B otany 105, 3 443-455 Thi s, P , Lacom be, T, and Thom as, M R , 2006, ' Hi stori calori gi ns and geneti c di versi ty of w i ne grapes' , Trends i n G eneti cs 22, 9: 511-519 V avi l ov, N I, 1926, S tudi es on the ori gi n of cul ti vated pl ants, Leni ngrad V oui l l am oz, J F, G rando, M S , E rgl , A , A gaogl u, S , Tevzadze, G , M eredi th, C P , and M cG overn, P E , 2004, ' Is Transcaucasi a the cradl e of vi ti cul ture? D N A m i ght provi de an answ er' ,i n Vi ei ra, A ( ed) , III S ym posi um of the Internati onalA ssoci ati on of H i story and Ci vi l i zati on of the V i ne and the W i ne, C entro de E studos de H i stri a do A tl nti co, Funchal ( M adei ra) , O ctober 5 - 8, 2003, pp 277-290 Zohary, D , and H opf, M , 2000, D om esti cati on of pl ants i n the O l d W orl d, O xford U ni versi ty P ress Inc. , N ew Y ork