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The Archaeology of Ajaṇṭā Author(s): Walter M. Spink Reviewed work(s): Source: Ars Orientalis, Vol. 21 (1991), pp. 67-94 Published by: Freer Gallery of Art, The Smithsonian Institution and Department of the History of Art, University of Michigan Stable URL: http://www.jstor.org/stable/4629414 . Accessed: 25/09/2012 03:19
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Stratigraphy of Overview Ajan.ta's
THE AJAN.A CAVES, CUT INTO THE CURVING SCARP OF A DEEP B.C., it is the later excavations that are particularly famous. Nearly all scholars agree that work on these began during the reign of the Vakataka emperor HariFeqa sometime during the latter half of the fifth century A.D.;traditionalists have long argued that the cutting and painting of the extensive cave complex then continued for some two centuries or more. By contrast, I have tried to prove that this great later phase of work at Ajarnta represents a sudden
gorge in western India's Deccan plateau (fig. 1), have long been renowned because of the beauty of their miraculously preserved paintings, all dating from the period when this Buddhist monastic site flourished over fifteen hundred years ago. Although impressive excavations had already been made at Aja%taby 100
18 ~20
Scale of 110 0 19
FIG. 1. General Plan of the Ajan.taCaves. Afterj. Burgess, "Report on the Buddhist Cave Temples,' A.S. WI 4 (London, 1883), pi. xiv.
TABu~ VAkd. white line=making of main Buddha. wealthy "owner of the cave": active patron 462-68 and 475-78. Cave 16: Varahadeva. from blocking out to painting *Specific dates given are only close approximations. 2). Cave 4: Mathuradasa. 2mU.totkacha Aurangabadl1 Aurangab. 18. no artistic r- 462 Cave AjauLta 463 0 1*m mE E .ad 4aEUE mm - ni *m -E MEn ** * Bagh (Caves1-9 - - Eu. Caves 17. For a detailed review of the evidence for this short chronology. a monk "attached in friendship [to the minister of Asmaka] through many successive births": active patron from 462 to mid-468 and 475-78.ad3 Aurangab. see "TheVakataka'sFlowering and Fall" (referenced in n.. probably 29: King of Rishika (named Upendragupta?): rules over Ajanta region until defeated by AAmakasbetween 472 and 474. 8 11mmm 14 15 16 17 18 * -UE mmmE m iim*** ~U ** 19 20EUE m mm 21 22 23 24 25 E ** H 26 26 RW 26 LW27 28 29 Gha. *: :: - more vigorous activity less vigorous activity intrusive work KnownMajorPatrons:Ajanta's VakTztaka Phase (A. Cave 26 complex (including Caves 25 and 27): Buddhabhadra. a margin of at least a year or two should be allowed.. 19. . SPINK 1.D.68 WALTERM. 462-80) Cave 1: The Vakataka emperor Hariseqa: rules 460-77. Harishena's Chief Minister: patronage continues until 478.E 3 4 7m.taka Patronage at Ajanta and Related Sites (462-80)* Rishlka feudatory controls region Recession caused by Rishika-AAmaka dissension 464 465 466 467 468 469 470 471 I Patronage beins AMmaka feudatory controls region Anxious Consolidation Period Limited Hariserja's Death New Florescence 4746474849404140 of Disrption residence. 20.
468. like its remarkable-if fragile and imperfect-state of preservation.THE ARCHAEOLOGYOF AJANTA 69 burgeoning of pious activity under the sponsorship of Hari$eqa and the elite of his empire during a period of scarcely more than a decade and a half (ca. and the fate that befell both him and the empire that he had gathered. prior to this time. Although the Asmakas' Cave 26 complex was among the most ambitious of the new caves sponsored at the site. with his close connections to the Asmaka court. the site quickly flourished again. with a unique record of this classic epoch at the very moment of its fullest blossoming. we can assume. At that point (our Level 1). But the world had surely changed. the local king. was able to continue his cave in a more or less normal way. Much of this history of the site will reflect the aggression of the Asmakas. The fact that. who had previously indulged in a lavish piety. while the caves of the local king were left unfinished and even (in the case of his sumptuous chaitya hall) quite intentionally abandoned. are part of the finally tragic story that Ajanta tells when we probe into it layer by layer. and in 471 (our Level 2) work was abruptly cut off. The entrance of the great emperor Harisena onto the stage of Indian history for his all-too-brief hour. even at this site far off from the capital. Suddenly changing his priorities from spiritual to military. their relations with the local feudatory king of the Ajarnta region (ancient Rishika) had soured by ca. the donation of his overlord. Although there is good reason to believe that none of these intrusive donations postdates the year 480. not one of Ajanta's excavators had ever faced a cliff before. abandoned their overall excavation programs. Then. During the brief Hiatus that followed-when. During the following year. only enhances the achievement. or the preparations for war now were choking the site off from its accustomed sources of support. From the moment of the old site's reinvigoration by a consortium of the emperor's richest and most powerful courtiers until its traumatic ending. he ordered all work at the site halted. and the skilled artists they hired. not satisfied with having usurped the Ajanta region itself. this main phase of work at Ajanta was all accomplished by ca. either for the better or (more often) for the worse. in the heyday of its power under the emperor Hariser)a. had hardly a rival in the world in terms of either its political or its artistic achievements. signficantly. or the rumors ofwar. except for that on his own caves and upon the splendid Cave 1. 480. Asmaka was battling for control of the Rishika region-no work was done whatsoever. feudatory to the Vakatakas. But Ajarna's aesthetic and technical virtues. even on these privileged royal caves. the majority of the patrons at the site. must have started to face up to reality. when patronage activity reappeared. the Asmakas had clearly assumed control of the area. who were ruling over the territory just to the south when Ajanta's renaissance began. Such energy combined with such quality-evident as much in the site's sculpture and architecture as in its paintings-would itself qualify Ajanta's Vakataka renaissance for greatness. Harisena and his courtiers. Most of these eleventh-hour donors were the monks still living there. 462-77).' Although so surprisingly little known. but not for long. the emperor Hariseqa. The next couple of years (479-80. But in 478 (our Level 5) all of the involvements of such original patrons ended completely. Only the monk Buddhabhadra. if I have read the evidence of his cave correctly. Thus begins what I call the "Recession"-a drastic halting of the site's patronage. in 475 (our Level 3). surely because the aggressive Asmakas were already threatening the region. By the early 480s. as if in a state of shock. what I call the "Period of Disruption") were years of frenzied and helter-skelter activity by all who could manage to add intrusive images in or upon or around the caves that the original elitist patrons had been so suddenly forced to abandon. a number of monks may have continued living in some of the caves for another decade or two. the turbulence ofAjanta's development constantly leaves its revealing traces in the rock. after less than two decades of artistic activity. have thus provided us. the aggressive Asmakas. work was now vigorously resumed on their own cave complex. In 477 (our Level 4) the great Harisena died-very suddenly. are the more significant because the flowering of the site under the Vakatakas came at the very height of India's Golden Age. The various "levels" or "breaks"in the site's development (see table 1) that I explore below can all be explained as revealing stages in the history of Ajanta's patronage. the Vakataka dynasty. anxious to get the benefit of such donations while they could and using whatever artists had not yet fled the declining site in search of more likely work. for no such caves had been cut in India for three hundred years. War. The Mmaka threat to the site's security only increased. Under this new Asmaka control. Even including the quickly fatal years of the site's disruption. rushing their images to completion in order to make such merit as they could. took advantage of the ineptitude of Hariserla's ill-fated son and successor to foment a widespread insurrection against the . when some of these same founders were to be counted among the site's destroyers.
Bagh. every carved or painted image. we can weave the complex tapestry of the site's dramatic history. They are. we can properly postulate their contemporaneity and often that of their contexts as well. divisions are arbitrary. or vanishes over time. Thus. when these features are sufficient in number and significance. every cave or portion of a cave."each containing. they also caused the dark age that so ominously shadowed their warlike path. and epigraphic factors.D. that Ajanta started its many centuries of stony sleep. because the upper and lower limits are so close in time. studying the way each separate feature emerges. cannot be properly developed in any other way. as it were. 2) are unique in this regard. in the end the history of the times depends upon the evidence of the cavesjust as much as the history of the caves depends upon that of the times. starting in ca. 3. I want to discuss not annual but rather event-determined levels. and other factors. As noted above. a curious fact emerges: whereas these outer records are necessary to put Ajanta and the other Vakataka sites into their general cultural context. even to believe. it breaks into a number of distinct levels or strata. just as in a "normal" dig. deprived of praise and nourishment. Instead I will concentrate solely on main shrine Buddhas. Ajarta and its associated sites (fig. is changed. SPINK imperial house itself.2And whereas the pace of developments within the sequence may vary slightly from what I have suggested. the most minutely. This is not to suggest that such discrete probings can be done at other sites. but they are not ve?y For reasons that I have explained at length elsewhere. by all counts. these arbitrary. we can then go to the written record-the inscriptions and the narratives of these times-in order to bring our abstract developmental sequence to life and into the life of those times. Indeed. That is. in the stratum under investigation. economic. Dharasiva. A. analyzable sites in the world. and because of the many revealing connections with known historical events. ideas. But we must make a vertical as well as a horizontal site survey. Then. however. Because the excavations of the Vakataka phase at Ajanta. at a given point in its development.70 WALTERM. and Banaoti are so abundantly rich in forms and features. painted. Finally. and 4A). which are not only of general interest to us but were of . And this fact is the more remarkable because Ajanta was created at such a crucial moment in India's cultural and political development. Ajarta's Vakataka development is quite discontinuous. the Ghatotkacha vih&ra. For our purposes here. 462 and ending in ca." As is the case in most digs. And though this history will still be mute. is year by year. Aurangabad (Caves 1. Such ruptures of course occur at the end of each of these levels. Their success caused the total destruction of the Vakatakas. and inspiration. it turns out that these records-long accused of being compounded of myth and exaggeration-are almost impossible to interpret. 480. which could include a host of architectural. will share certain distinctive features with its counterparts elsewhere at the site. evolves. as well as the most totally. which must be radically transformed by what the caves reveal. sculptural. and technological features spreading horizontally throughout the caves. until they have been tested against the undeniably solid evidence of the caves themselves. As we probe into each level-the eye rather than the shovel being our tool-we find directly comparable morphological. There is no question of a report on all the features that one must consider. neither the starting date nor the ending date can vary by more than ayear or two. and to read its story. technological. focusing upon the five very distinct points of rupture that can be found in the overall development of the site. using slightly elastic bands for our woof and fibers from the date-palm for our warp. But because they themselves could not manage to hold the empire together. when we consider all of the features together."it should be possible to speak of an "archaeology of Ajanta. we should be able to define and describe a series of vertically ordered planes or "levels. for it is there that the drastic temporal breaks occurred.D. the features belonging to that level of historical development. Thus. Admittedly. It was within that darkness. it both describes and explains that classic moment in India's past from which later generations in India and beyond continued to draw forms. Yet when we putwhatwe have observed at the site together with what we know from written sources. our understanding of the history of the times. As a richly informative illustrated history. only a few shards from our dig can be shown here. the most practical and informative way to describe the development of the site. iconographic. GeneralConsiderations Since Ajanta is a "dig. this variation too must be slight indeed. leaving a spectrum of unfinished or just-finished forms and features embedded. as in an archaeological dig. reflecting the way that its patronage was affected by political. Needless to say. I have divided the Vakataka developments at these cave sites into nineteen separate annual levels. A.
Plans of shrines at AjanA and some of its associated sites . 2.THE ARCHAEOLOGYOF AJANA 71 Cave 1 Cave 2 Cave 4 ~ Cave Upper 6 ~ Cave Lower 6 Cave 7 Cave 11 Cave 15 Cave 16 Cave 17 Cave 19 Cave 20 ~~IRhr7~2~ | Cave 22 0 l54 S Cave26LW i! F : 26 XRC Cave 21 Cave 26 Cave 27 Ghatotkacha vihara Aurangabad Cave 3 FIG.
the "biography" of any given example might span much of the site's development.' Nor were the other five immune from pain.4 Two others (in Caves 26 and 26 LW).72 WALTERM. SPINK overwhelming interest to the site's patrons. like so many of its Ajanta counterparts. just as each new level always begins on January 1. with the spaces between the dates being somewhat reduced or expanded as a consequence. the plastering. were we considering windows or doorways. no otherimagesyet reachedin courseof excavation. all but five had to be rushed to completion at one or another of the different "break points" in the site's history. I shall ask the reader to refer to my previous articles for a more detailed discussion. 7. the carved decoration. Rishika was at peace.D. like some of the other twelve. after one took into account the original conception. for this will not affect the underlying sequential connections.8 Indeed. the painting. I shall also briefly mention the main Buddha in the Ghatotkacha vihara (donated by the Vakataka minister Varahadeva. 460. The single (not particularly impressive) image at Ajanta that seems to have survived unscathed by the site's turbulent history is that in the left wing of Cave 26. 15 all rushedto completion. 11.5 Finally. other than those caused by traumas over which the patrons had no control. Two of them-indeed the most important of them all (in Caves land 19)-were never used as intended. the proper shaping. even drastic revisions. but even that image enjoyed no more than a few months of what one might call decent worship. whereas the Aurangabad Buddhas. along with the obsessively pious local king and a number of other wealthy and powerful devotees. it is remarkable that of the seventeen main images finished at Ajanta during the Vakataka phase. Or it can be stretched a little in one part of its composition and shrunk a little in another. which. were probably donated by the Asmakas. whose territories lay just to the south. Furthermore-and this is crises came. while under the authoritative rule of the new emperor. Thus. the great trade routes that passed through the region were open. were less hastily finished in that same year. upon which attention was not so compulsively focused.7 The most ambitious of these-the chaityacomplex involving Cave 26 and its four wings-was sponsored by a Rajneesh-type monk named Buddhabhadra. but they should not be thought of as absolute." as his donative record proclaims. although all of the seventeen images atAjanta thatwe shall discuss were invariably and even compulsively brought to completion. when they decided to reinaugurate patronage at the famous old monastic site in the spectacular ravine formed by the headwaters of the Waghora river. Having only recently defeated their old rivals. Level1: Early462-Late 468 Buddhas of CavesLower6. The Ajarta region was among the extensive feudatory domains inherited by the new Vakataka emperor Hariseqa when he succeeded to power in about A. patrons were most significant-when always concerned to finish the Buddha image. Considering how little we know about the biographies of most Buddha images in India and how seldom we can explain the problems affecting them. Mmaka. suffered unanticipated compositional and/or iconographic adjustments. when traumas disrupted their development. 26 roughed out. Instead. even certain reshapings and refittings. stupas to contain Buddhas of Caves 19. to the exclusion of almost everything else.6 The dates that I shall supply are from the neatly framed tapestry I spoke of before. For this reason. being under Mmaka sponsorship. Buddhabhadra's power and wealth can be explained by the fact that he was the closest friend of the great minister of Asmaka and had been so "throughout many previous existences. was rushed to completion in 478. like the Cave 26 complex at Ajanta. Because they are products of precisely the same patronage. who had intimate links to the court of the powerful neighboring province. in the interests of brevity. Nor will it affect the remarkable fact that all of our break points happen to occur on December 31 of the year in question. the Cave . Within the first five or six years twenty-four major excavations were already underway. the roughing out. Such factors surely influenced the chief minister of the empire. The Ghatotkacha Buddha. By contrast. a rush to complete these main Buddha images generally characterizes the "break point" of each level. it can always be stretched a little or shrunk a little. I shall not of course be able to discuss all of the characteristic contextual materials that define each of these levels nor to report on the whole dig. I will list here the various images that had eitherjust been hurriedly completed or (in a few cases) hadjust barely been completed in normal course when each level ended. work once started generally proceeded upon such images with no breaks at all. the fitting. this was not true of a single one of the caves in which they were located. which extend out beyond the images with an even more poignant story to tell. Because of the way the cloth was made. who also sponsored Cave 16) as well as the main Buddhas in Aurangabad Caves 3 and 4A. the Mmakas.
= 3'"'_ J9 S - S U x _~~~~~~~~~~~~~~~~~~~~ _~~~~~~~~~~~~~~~~~~~~~~~ JGE --- z R . TA 73 26 complex was being built in honor of this very official.i . revisedlater in 468..10 This is clear from the forms being excavated at the time of the break (many of which can still be seen). for they are very different from those produced when work was resumed on most of these caves over half a decade later... it is at this same developmental level that work broke off on nearly every cave then underway at the site. 3). Quite possibly the earliest conception of all is that of Cave 11 (fig.5z ~.. only four Buddha images were underway in the site's viharas.. and all display certain significant early characteristics not to be seen again in the images that were later developed at the site... work on it suddenly broke off.*.. and its importance (mirroring the importance of its patrons) is attested to not only by its size but by its particularly sacred function. s~~~~~~~~~~~~ . Main Buddha... at this very time. AAAUM 412/69. P . However. where the seated Buddha is backed by (or revealed from) a stuipa.~~~~~~ FIG. Caves 19 and 26.. Cave11."1All of these images were apparently intended to be located in the center of the shrine chamber. 3. when the great central hall of Cave 26 had scarcely been roughed out and its stiupa barely revealed.THE ARCHAEOLOGYOFAJAN. *>2 U t S'. A = . suggesting that the first image type conceived for a vihara at the site was directly derived from the types being created in the site's new chaitya halls.9 Significantly. Begun 466.. . Since the F . At this moment. . completed 468.
The fact that both were redesigned and refurbished a decade later-the time to which both their throne backs and the throne base of Cave 15 well suggest that they remained unbelong-could painted and undedicated at this early stage in their varied careers. As we might expect. not only in 468 but for the whole next decade. completed 468. AAAUM 41/68. revised 477 78I AAAUM 1569/73 . their shrine doorways had not been hung at this time. However. Indeed. if they were finished at all (fig. Main Buddha. 7). the focus in these viharas turned out to be on the image alone rather than on the stiupa or any ritual involving its circumambulation. In Cave Lower 6. Had there been any serious interest in the stiupa in Cave 11.although the large size of the halo. but in the rush of the moment this was never done. characteristic of all these first four images. 4. the assumption that the chaityahall image was the source for that in the viharas seems very compelling. Main Buddha. the contemporaneous image (fig. theywere finished most expediently. Begun 468. 4) is already free of any link with a stuipa.74 WALTERM. _ I Cave Lower 6. the floor around it would certainly have been finished rather than left completely rough. Begun 467. FIG. 5. Cave 15. 5-6) were presumably originally intended to be centered in the shrine-a convention that continued even into the 470s in Cave 17 and in the original plan for Cave 1. as originally planned. 4. SPINK FIG. The Buddha images in both Cave 15 and Cave 7 (figs. completed 468. had no shrines or images whatsoever. earliest Vakataka viharas at the site. as Cave 11 itself shows. can perhaps be explained as a vestigial connection: in Cave 11 it traces the outline of the attached stuipa's dome.
*._ iI-F ...._ _-sffi __ I 1 _Sz_ _s__ ^--t_-__ $. _. ill <_ _fi..X t! i 1|1 | |l ' _1 w-d ? X S__f J 1 l l I _^ _ | '. b E ._lFss 1 | s_ _>ilCiL l 111 1 $ 11 h i Ki isl | S.4. :_1i ..R.] m l | -'baN_ _S. < . 1 v * s +' F-8f: tili.* FIG.. .W._Bu_ . completed 468.. . Begun 468.. ....e.. .. _2 . .Yks.bll.... i i li _ : 1. . ' ev ..S }... tt ?2 t_t i * r....=f SBSt i I __ ex. ^12 C.b"' 1. v e " x .:. gtes...__ e. X -'' vev .... .... 1 z _. Ws' " S-||>. 1|_ __ _51 ?|11 | _mSV " 4e ...S El m | _ = = W..: : .l[ 1i sZ Wa Es. " .'::+...7 e in2. 4 * .. --?'w.. :*.+... iWsi . *' is ^ -' ........e : iAR.O t ... . > ' t. . w>p-is j>.t .THEARGHAEOLOGYOF AJASTA -Y 75 _1 _ t... 6.. IEZ . . .. -. AAAUM1446/73.. i. _ .. w gS __ t t @ 11 S | :_ 1 S111 l 11!1 E | .. Sketch showingCave7 image in 468. _F?iX_. SsNi x.. .... \... lug 111|11 s...j>#.Cave7..- ?....s _f 1 _3RF Hl=e.s i..x _>. * */..*.......^. !.. b Z r . >5 . t .b.?s ..r * -.. Main Buddha... - . ... ...w. F. j:..... t w jLl &. .....W11 i . *. N .. 7. LE v S ---AF .e =. t shtsbl-tD\s s . ..E s_.Ye zR_..<Aiiil 4' E * . .?n B .. _= _ - 5|_S. 11s u ..lf _ : ..| 2ailC'> _ *5.Xt. S .' _111 -- ?l * _j ... . ' E__ __ _ _i_ __ _ _S _F _ So s Ms_ o' ' ' 'n FIG...<?e S t lrh > *io ms-S 0 $ W ' t. | iKiq ... ..lGl [4Jat '' ' C. . .."w.... . E11 11 -_ _t.. _Wi _ iNF$#e _ _ w * 3 _ _2__ ! B !fl5 ... ..v .. ._.-.0. !111lii3bEe ffltgx-a 'sJ3rg -e k 11 ilE 1 e NliltX 1M k05iti iF X iIXv l-Nil i-^ ...... revised477-79.miE__ -!c_ _.Fw . eX .j... E < > \-W l1 T S..\ ' ? " .i a - .
the only one of the four where the shrine was plastered and painted at this point.76 WALTERM. Cave 20. on or under the vyalas. 7.12 I have called this break the "Recession" of 468. for although the craftsmen had penetrated the shrine antechamber before they were laid off. who was-not surprisingly-largely exempt from the sumptuary cutbacks that afflicted lesser figures and so was able to continue the overall course of work on his Cave 16 during this critical period. Main Buddha. a serious but notyet desperate political or economic problem led to these cutbacks. At ~A?- FIG. and 15 proves. the careful carving and painting of the previous years now gave way. . of attendant bodhisattvas.118. the patrons were discouraged or perhaps even forbidden from making similar efforts.13 In any case. even in Cave Lower 6. in all four. it was barely half completed at this time. This is probably why the rear half of the simply painted ceiling and the shrine's rear wall were never painted. it was not a total and traumatic break. at this early date. Begun 470. the intended image still lay behind some twelve feet of solid rock when work was given up. and it surely explains why the decoration of the left wall was never finished. for instance. clear evidence indicates that there was no time to spare. as well as a characteristically early throne base. Furthermore. It does. which had certainly not been anticipated even a year earlier when the site was still expanding so exuberantly. for work still continued both on the splendid cave complex (Caves 17-20) of the local king and on the sumptuous Cave 1. during 468. Also. 8. it finally all came to naught. for time so quickly ran out. Apparently. As for the cave's program of painted decoration. SPINK What distinguishes these images from later ones is the absence. there was clearly time even if only a few months-for those patrons to bring their images. 11. But even in this powerful minister's cave. The only exception involved the Vakataka's chief minister himself. however. or barely finished. where it shows (as in Caves Lower 6 and 11). quite expediently. no dwarves frolic. to or at least toward completion. to efforts that show every sign of haste. AAAUM15. completed 471. the halo is much larger than it is in a typically late image. and. appear to be the case that where the shrine areas had not even been reached in the course of excavation. the throne back is far simpler than later on. which the emperor HariFeqa himself had only recently started. in which lions appear under (but not-as later-supporting) the throne and deer flank a simple wheel. And all of these images are of course unfinished. even though they were fully plastered. as the evidence of the images in Caves L6. In its effect on the site.
8-9). the shrine was redesigned to save time in its excavation. in the second level. it is clear that Upendragupta's pious goals-enthusiastically announced in his Cave 17 inscription as "filling the whole world with stupas and viharas"-now had to suffer certain practical limitations. drastic cutbacks were now ordered.c iS Sg x | #X_l3?1v. FME_e li 1 2 C | SiE ! l j i1771 | v@ 4 w | a |~~~~~~~~~~~~~~~~~~FG Buda Cave vie rt Begun1 9 Mai 20 cml '''eted 471 t | I~~~~~~~~~~~~~~~40 Hningo R318 . before an explanation ofwhatwas happening will emerge. we have two more levels to go. Buddha image in Cave 19finished but not significantly blocked out. whose Cave 1 progressed with great care and creativity during this period. Finally. At least this is what its many forms and features suggest. used. and of his overlord Hariaera.THE ARCHAEOLOGYOF AJAN. it was under direct Vakataka control and apparently exempt from whatever problems were afflicting Rishika at this time. and expediency rather than aesthetics became more and more the rule. from 468 onward. The two rear cells were omitted even after matrix had been reserved for their elaborate door frames. while its fine shrine doorway was hastily plastered and painted _ 4. Buddha image in Cave I left merely It is of course significant that. hoping to find work on the new Vakataka caves at Bagh-a site that may have been recently begun because. with one of Harisera's sons as the viceroy at nearby Mahismati. numerous halffinished or half-started motifs were masked with mud plaster. so many unemployed and already experienced excavatorswere suddenly available. Level2: Early469-Late 471 Buddha images in Caves 17 and 20 rushedto completion. In fact. and a drastic abandonment of all activity in between. was finished off with mud.TA 77 this point in our excavations we can have no valid clue as to the cause of this Recession. but despite the splendid quality of the work that is generally maintained throughout his feudatory's similarly "privileged" cave complex. The emperor Harisera's Cave 1 progressed with an unhurried authority during this period. the beautiful Cave 20 image (figs.14 Furthermore. It is clear that the very best of the workmen had been selected to work on this impressive excavation and on Upendragupta's equally superb Cave 17-20 complex. the local king. for dark clouds were gathering on the horizon. work at the site continued only on the benefactions of Upendragupta. Especially in the beautiful little Cave 20. not quite yet fully carved. Many of those not so chosen probably went north to ancient Anupa at this point. in desperation.
but nothing more was done on the splendid central Cave 17-21 complex. atAjarztais very clear. there is not a trace of the grime that darkens the painted surfaces in caves used constantly for even a year or two.78 WALTERM. for by 475 he was clearly no longer a presence at the site.15 In Cave 17. which had been sponsored by Upendragupta. but a number of significant features in the cave were never finished. The most significant event connected with the Hiatus-in fact the event that caused the Hiatus-was the conquest of the region by the aggressive Asmakas. 11). We should of course say the formerlocal king.at thesitedunringthis period.Many craftsmenfind at employment Bgagh. a brief record of the king's donation was all too hurriedly recorded. Main Buddha. or Hiatus. On the other hand. "having expended abundant (wealth)" on sacred monuments but all too little on his arsenals. as I believe. the local king. the main phase of patronage at the site. Just as in the closely related Cave 20. had been intended as the ceremonial center for the whole site. Upendragupta's beautiful chaitya hall was never put into use. Similar problems afflicted Upendragupta's Cave 19. although the Buddha image itself was successfully completed (fig. lasted only some seventeen years (462 through 478). such an inauspicious omission further shows how quickly time ran out when Upendragupta was losing his control of the region-a conclusion confirmed by a number of lesser unfinished details in the court. was never placed in the panel planned for it. even though still far from fully carved. he had been too pious for his kingdom's good.eitherproductiveor destructive. the long dedicatory encomium. 10. AAAUM 512/77. there are other indications too that work broke off on the cave. now that the Mmakas were in control. SPINK finished without a problem (fig."9 Not surprisingly. on the left porch pilaster.Strict administrative control of site continues. which. the beautiful chaitya hall. Although it is evident that the image was . Perhaps.16 Most significantly. Hiatus: Early472-Late 474 No patronage activity whatsoever. did indeed get applied. where the same thing happened. and that it was in fact abandoned. The evidence for a total interruption of patronage activity. The fact that a number of the expected symmetrically placed garland hooks were never put in place suggests that the interior was never properly fitted out. the shrine walls were only plastered but never painted. some events that occurred during the Hiatus seem to require it to have occupied a year or two at the very least. it appears that the new rulers specifically disallowed all worship there. 10). Outside. Here the wide panel prepared for the inscription remains poignantly empty. which must include the Hiatus. Cave 17. before it had even been used in any consistent way for worship. completed 471. Indeed. which was being built in honor of the minister of Asmaka. It cannot have lasted very long if. even though the span of time assigned to it is necessarily a guess. in better days. intended to mirror the famous nagaraja on the other (west) side.just as they never allowed any more work to be done on Upendragupta's Caves 17 and 20. Begun 469."8In the courtyard the yaksha (or yaksh-). for it could house forty monks and was one of the few FIG. it is clear that time ran out. like anotherJustinian. which by good fortune had already been composed and sent down from the capital. work was vigorously renewed on the Cave 26 complex. In 475.17 while the paintings inside the cave remain suspiciously clean. even though for purely practical reasons Cave 17 continued in use.
when the AMmakas had given up their authority over the site. which Upendragupta had selected as the site's focal image. All such evidence finally clarifies what camne before. . just before the Hiatus. the Mmakas may have gone further than this. In any case.21 Then it returned assertively.THE ARCHAEOLOGYOFAJANTA 79 caves available for occupancy (and indeed already occupied) when the Asmakas took over the site. as if released from bondage. were already gathering over the Ajaqta region at that time. It explains the regional political and economic problems that caused the Recession of 468. this particular image never appeared again until 479. which lay beyond. AAAUM143/68. It even explains Upendragupta's touchingly talismanic boast. after having appeared frequently on doorways in Upendragupta's caves in the early 470s. Perhaps the AMmakas that this hieratic image type was felt too closely identified with their old rival's power.20But as for the cells attached to Upendragupta's chaitya hall. made in the Cave 17 inscriptionjust when the Asmaka aggressors were beating down the door. for it seems clear (given the hindsight available after the Hiatus) that the dark clouds of Asmaka aggression. the Mmakas brutally cut a passageway right through them to create a more convenient approach to their own Cave 26 complex. Indeed. 11. It appears that they even banned the use of the standing Buddhawith both anns raised. which would ultimatelyspread so disastrously over the whole of central India. Main Buiddha. Cave19. And it explains the surprising cutbacks in Upendragupta's lavish patronageparticularly in Cave 20-as the reality of war began to loom. Begun 467. completed 470. that when he and his brother were ruling w *i 4 t e? - ~~~~~J JIPoL <:r FIG.
____ ____ ____ ____ _____ first phase of work on imnage t second phase of work on image ... etc. no throne legs and no lions Base with lion throne legs. Cave U6) Wheel absent (pralambapaidasana) Nas at base (pralambapadasana) Wheel present (pralambapadasana) Wheel fused with lotus shaft Wheel raised on lotus shaft Base with nubs at corners (pralambapadasana) Base with nubs at corners (padmasana) Base shows small robe end on seat cushion Base with arcing doth over front (padmasana) Base with scalloped cloth over front (padmasana) Makara empty mouth Makara dwarf emerging from mouth Makara rests paws on throne arm Makara frond or bird's head from mouth Makara dwarves as well as the latter Seated figure over makara No figure over makara Running or soaring figure(s) over makara Multiple figures over makanx. Vyalk omitted Vyalawith no dwarf attendants Vyalawith one dwarf on or under Vyalawith two or more dwarves on or under Vyalawith beads from paws as well as snout Pillared throne back Pilasters frame Buddha panel Flying dwarves above image Flying couples above image Both flying couples and dwarves above image Bodhisattva attendants Standing Buddhas as attendants Attendants on lotus pedestals Attendants with halo Avatokitekvara with long-stemmed lotus Attendants * 11 7* 15* 468 469 17 470 20 1* It 4 471 Hiatus 475 476 e on long-stemmed lotus pedestals . SPINK TABiL Introduction of Various Buddha Features at Ajaria and Related Sites 2. with/without devotees Base with dwarf throne legs instead of lion legs Wheel plain (with decorated hubs) Wheel festooned (e. Cave 1) Wheel garlanded (e. (Images with listed features were carved no earlier than date indicated) 2 21 7t U6 15t 26 22 26RW 26LW all intusive 16 Aur3 Aur4A ima es 477 478 479 480 isages carvedat apprxmatly thes date: L6 467 Buddha: Padmasana Buddha& Pralambapdasana Buddha placed in center of shrine Buddha attached to rear wall of shrine Buddha& unusually large halo Buddha: smaller (usual size) halo Buddha: hands proper right in abhayamudra Buddha: hands in dhavmacakra mudra Base with lions but no throne legs Base with throne legs but no lions Base with devotees. dancing.80 WALTERM.g.g.
10). work started up very vigorously. How strong and how effective the administrative control was during this troubled time becomes clear when we contrast it to the total anarchy that reigned during the Period of Disruption. FIG."22 Evidence that the Hiatus was a distinct interval separating the period of Rishika control from that of Asmaka control can be seen in the decisive replacement of older (and suddenly outmoded) forms by new and better ones. the burdensome double stone projections used for cell doorway fittings from 470 until the time of the Hiatus were expediently cut away when the sensible new recessed mode appeared in 475. completion themain imagesand theirimmediate of After the Hiatus.whichwerereactivated nearly all of the abandonedcaves and initiated in somenew ones startingin 475. at least in part. along with the thickening of the walls in which the doors appear. which would suggest a break in time long enough for such ideas to reach the site or to be developed there. right after the Hiatus. At the same time new window and doorway formats and a number of decorative elements were similarly "imported" at the very beginning of this second "heyday.)23 Evidently. Thus. 12) is a case in point. Work in programs. was less than thorough. it had apparently been roughed out by the time of the Hiatus to the extent that deep cuts had been made along either side. to the degree possible in caves so generally incomplete. Modeled. can be followed at Ajanta almost year by year (see table 2). so the telling evidence remains. in Cave 1. 4~~~~4 Level3: Early475-Late 477 A A Cave I Buddha imagefinishedbut neverdedicatEmperor's ed orused. who could find no work there-as it does its continued occupation and the preservation of the insistent exclusiveness that had characterized the site's patronage from the start." Not a single image of any type whatsoever was added to any of the caves at the site during the Hiatus. including the emperor's Cave 1. the cutting away of the old forms. by and large. Then. these preexisting cuts were converted into (necessarily pointless) "corridors. as we might expect. Immediately thereafter (in ourLevel4) attentionis turnedexclusivelyto thehumed contexts. continued 475--77. they had "subjugated prosperous countries such as AMmaka. in all cavesexceptthe Cave 26 complx. (Fortunately. aresuddenlygiven up at theend of 477. and worship must have continued too. on the Asmakas' long-abandoned chaityaCave 26 complex. with a central placement planned. when the major patrons had lost their rights and privileges and anyone could put any image almost anywhere. as well as on various other major undertakings. on the Cave 17 image (fig. most of these too appear to have come down from Bagh. when work on the image was taken up again after the Hiatus and a central placement for such images was no longer desired. Thus. AAAUM11/68 . between the time that work on Level 2 broke off in 471 and started up again on Level 3 in 475. the new fitting mode. where the sandstone is so friable that projecting fittings were not practical. Main Buddha. distinct changes were made in the format of certain images or elements. Finally. Begun 471."which started in 475 with the Mmaka conquest. Cave1.24 This does not so much suggest the site's abandonment during this period-except of course by the craftsmen. The placement of the Cave 1 Buddha (fig. which were in low-priority locations. had been developed at Bagh. in the case of Buddha images. 12. The fact of a strong administrative control over the site of course explains the generally orderlyway in which it grew and may also explain. where the very softness of the stone probably encouraged such elaborations. the remarkably consistent changes in iconography (as opposed to style) which.THE ARCHAEOLOGYOFAJANTA 81 happily together some years before. monks continued living at the site in this fallow period.
and a shock wave surged through the site. Apparently. Several factors suggest that the cave was neither used nor dedicated: it was not inscribed-at least no inscription is extant. remained totally untouched by further excavation or decoration. no fewer than twenty-five caves were simultaneously underway. when new donors were scattering votive images throughout all of the caves that had been surely dedicated. despite such practical problems. 13). U6 (fig. The renewal of work on certain lesser caves was somewhat delayed. 17). 21 (fig. 7 (fig.27But itwas certainly not a wasting illness. in the Dasakumaracarita. the image shows not the slightest trace of smoky grime. 5). 26).82 WALTERM. almost certainly because at the very start of this new phase there would have been more work programs than workmen to work on them.AAAUM FIG. and the rest of the cave was was hardly moving nicely toward completion-there two weeks' more work to do. 4 (fig. when Level 3 concluded with the sudden death of the great emperor HarieKra. It is inconceivable that. MainBuddha. the image was peremptorily abandoned before being brought to life. We are told by Dar)1in. 18). 23). everything went on exuberantly now.totkacha and in Aurangabad Cave 3 and 4A (figs. the donations of the defeated local king.25 The image had just been finished. 22). .completed WG/14. Ten of these (excluding Cave 1. Similarly. then. the emperor would not have ordered it XR~~~~~~~~~~~~~~~~~~~O 478. or (considering the perverse machinations of the Mmakas) it may have been by poison or the knife. 14-16). no such intrusions were put in the splendid Cave 1. 26LW (fig. 6). SPINK Only Caves 17-20. in the Period of Disruption. 26RW (fig. 24). First of all the sumptuous Cave 1 (which I have argued elsewhere was HariFeqa's own donation) was never completed. whose image had just been completed) had images then underway or (in the case of Cave 26's right wing) about to be. By the end of 477. 21). and garlands were never hung from the hook provided in the shrine antechamber's ceiling medallion.26 HariaeTa's death was surely sudden. and that on the murals alone-when everything abruptly stopped. These are Caves 2 (fig. But then HariFeqa died. One could also include the images in the vihara at Gha. 15 (fig. 16 (fig. having sponsored the finest viharain the whole of India and standing to gain much glory from its completion and dedication. but the best evidence bearing upon his death-and perhaps even the manner of it-is embedded in the site itself. Begun 478.Cave26 RightWing. 26 (fig. that the death of the greatVakataka king brought on a time of great strife and presaged the fall of the empire. Nonetheless. It may have been from a heart attack or a stroke.13.
. FIG. 15 Main Buddha.Benares694/55.THE ARCHAEOLOGYOFAJANTX 83 14~~~~I zghdra FIG. Ghatotkacha 14. 16. AAAUM WC/28. 3-. Begun 476. Main Buddha.15. E:S-- 3 4A. Reproducedfrom AAAUMslide. Aurangabad FIG.Aurangabad 3.Aurangabad Begun 478. AIIS. F IG. completed 478. Main Buddha. Begun 477. completed 478. i W~~~~~~~~~~~~~~~~~r 5. MainBuddha. completed 478 (intrusionsadded 479-80).
... .. with respect only to images. In all of these other caves consistent excavation work is now halted completely.. Begun 477... at the very moment-the specific 'archaeological level"-thatwork breaks off in Cave 1.. . 4... and then only at the point when it was just about to be dedicated and put into use. this break precipitates a pervasive work cutback throughout the site (again with the exception of the Cave 26 complex). Suddenly. except. Thus... If the situation in Hari4eqa's own cave appears to support his sudden death.... importantly.... We shall meet these images again. finished if he had known that his death was imminent.W~~~~~77 - I _ ~ u .. the situation throughout the site does so even more dramatically...84 WALTER SPINK M.~~~~~~~~~~~~~~~~~~~....... AAAUM1011/68. at the end of our Level 4.. But... VN If w- !!|1|i t i t ~~~~~~~~e t < . FIG... all attention being anxiously directed to the suddenly crucial task of completing and dedicating each cave's main image-often in a most expedient manner. 17... significantly.C t^ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ .Cave26. completed 477... the break in our Level 3 involves the Cave 1 image alone. in which work shifts anxiously to the main Buddha images alone.. ... when the time allotted them has run out. scarcely a few months hence...... work is drastically affected in every other ongoing excavation program throughout the site. Main Buddha.X. in the AKmakas' Cave 26 complex.
.... 11)..XH _ i gwS I | * | ' i N1_ _ Z = !i LkA W t . ...ei_1 'iE_f_. Begun 477. E . whereas in the related panels carved during - - ? w ss'Wl : I h_ . '_ w _1|11 _||s!-_ _S S ...lu i' _ % _4'_ _ a . on the one hand.. this new pralambapadasana image does not properly fit the space allotted to it. _ 4 [Ad _i | b D k s | F ..I_1 '__S ... the chaityacave that was now surely intended to replace Upendragupta's rival Cave 19 as the ceremonial center for the site.. AAAUM1100/66. w _ _ - S . Consequently.. The now-necessary bodhisattvas had to be relegated to the wings (they can be found in miniature just beyond the frame....... 3 v t_r '.*E-t.._ . E i...Minfo | h ..... on the other..X: ''....... for the connections between the two regions were apparently friendly when the site was inaugurated. _ I ...... _ . and in the donation of the Vakataka's chief minister.: s_. Cave 16 (fig........"_ \ F......THE ARCHAEOLOGYOFAJANTA 85 Other than those thatwere rushed to completion in 468 or 471.. a newfangled type that was never carved at the site prior to its similarly late (ca. for its stupa had already been roughed out when work on the cave broke off a decade earlier at the time of the Recession. 18. W P _ ..r @ B - | 111 e .. (F 9 ... . completed 478. i]e iSIis .. __ _ w_ t R! 11.. on the stiupa drum).... MainBuddha.Cave16. 477) appearance in this cave.. 18)..... It surely was not intended to contain its present image. like that in Cave 19 (fig.. w=Xffi ....... 17).. S s1 L u l b I 1Sid1Y _ _ _ _ _ _ _ Ei | | | s _ _ S S S D - l | I | I __ | | __ | | I I I ___ __ __ lN | | | FIG... It has a complex history. the only image that may well have been fully carved (and quite possibly painted) by the time of Hariaexa's death in 477 was the main image in Cave 26 (fig. t: _' S# _ . r _ _. At that time it maywell have been intended to contain a standing image..
Panels unfinishedwhen workbroke off in 480. 19. and 22. where there are no such constraints. 7 (Phase 2). and in Aurangabad 4A completed normal course. FIG. The shock waves that ran through the site at the word of Harisera's death could only have been exacerbated by a knowledge of the incompetence of his son and successor Sarvasena III. Parinirvava (left) 478. not completed in wiiUlbe completed thePeriodofDisruption(479-80) butno longerin accordwith theirpatrons' original intentions.20. they are included comfortably. and they were probably a bit larger. ambulatory.which also breaksoff Jurther at abruptly theend of this level. 21. and dedicatedin time. while the group at the right had to be carved separately and then plugged into the floor. than their constrained counterparts. 15 (Phase 2). 4. Cave 26RW. 16.Buddha imagesof Cave U6.86 WALTERM.Buddha images of Cave 26LW. it was apparently not considered appropriate to cut the figures of these now-conventional devotees into the structural base of the stuipa Consequently the devotees at the left are carved from matrix that had happily remained on the floorthough a bit too far back-from the time of the cave's abandonment in 468. In the central stuipa. as are the kneeling devotees below. Largest Buddha (right) late 479. Aurangabad 3. and Ghatotkachavihara rushed to completion. SPINK the following year in the ambulatory (figs. AAAUM1183/78. What followed was a FIG. Cave26. AAAUM 1203/78. Padmasana Buddha (below) early 479. Buddhas (intrusive). Cave 26. along with workon the Cave 26 complex. too. Buddhas (intrusive). 19-20)." Level4: Early478-Late 478 Main Buddha imagesin Caves2. K . Buddha panels (above) 480. But at least the righthand group could be placed as far forward as desired. ambulatory.
A.THE AR(CHAEOLOGY AJAN.' AAM 5. in general. 7 and 21). and a certain amount of expedient FIG.:"y'. the main image (fig. they concentrated on the main Buddha images alone. In Cave 15.22.'. the patrons at Ajanta tried to continue with the painting programs.z:: FIG 2. t z 5P' Ws. were not touched at this point.00 s _~~~~~~~~~~~~~~~~~~~~~~ drastic halt in nearly all of the ongoing work programs at the site. along with the total decoration of the shrine. which had been left half carved in 468. Two of the six great standing Buddhasthose flanking the shrine doorway-were also rushed to completion at this time. Begun 475. 22) was quickly plastered and painted. In Cave 4. even though the shrine as a whole was still in the course of excavation. which lay close to the center of Mmaka power and were almost certainly supported by them as well. 21) was hurriedly completed in 478. MainBuddha. but thiswas done most expediently. but the other four had to be abandoned.ZTA OF 87 's.Cave4. the old image. which were already plastered and in normal course would have been painted prior to those of the shrine. ai Bdda.30 In Cave 2. the colossal image (fig.copleed47.. made in the Period of Disruption. Z: "'1~~~~~~~~~~~~~~~~~~~~~~~. and their very immediate contexts.'. 5).. was now hurriedly finished (fig. The only exceptions were the "Amaka" Cave 26 complex and the recently started caves atAurangabad. the shrine antechamber walls. ' a . then very hastily and informally inscribed. AAAUM78/78. all of the many Buddha images that cover it are new donations. mbw. ..Cav x~~~~~~~~~~~MO 2 Bgu 46. completed 478." Where they could (as in Caves 2. By contrast.
MainBuddha.88 WALTERM. havingalreadyrenounced his original excavation program by cutting his main image (fig. Cave22 provides a somewhat similar example: the original donor. suggesting the haste that was necessary. where the old image was updated startingin 477 (fig. the image in the main stiipa (fig. 24) was fully carved. But the image was plastered and painted.23. the images in the Cave 26 complexwere completedwithoutincident. Begun 477. the very new and up-todate pralambapMdsanaimage (fig. which in normal course would have been done first) was hastily decorated with contextually related paintings.31 By way of contrast. only in the Period of Disruption. In Cave 16. the image (fig. this was also the case in Cave 7. 23) was rushed to completion even before its hair curls and its conventional throne back motifs and attendant figures had been fully defined. even though the cave's rear wall (but not the ceiling. In Cave 21. finally had to share that area with other donors in order to get his image completed at all.for time ran out all too fast. 18) was fully completed. and a good start was made on the large standing Buddhas that were to attend it in the shrine antechamber. Indeed. As we have seen. SPINK paintingwas done in the shrine and shrine antechamber. reflecting no such anxieties. 25) in the rearwall of the half-completedshrine antechamber. in that same year. 6) but was only finished after the site's constraints had been broken and new donors had startedadding their own images to the cave. the Cave :- 'w'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~' FIG. . completed 478. the Buddha image (fig. 17) had already been completed. perhaps even before 478. along with those in the shrine antechamber. In Cave Upper 6. AAAUM1389/73. butits context-the splendid pavilion inwhich it is now placed-was left totally unplastered and unpainted. even though time ran out before the shrine door could be hung.Cave21.
Cv EW'. _ . ?'# rA::. _~~~~~~~~~~~~~~~~~~L opee 7.THE ARCHAEOLOGY OF AJANTX 89 ! * FiG 24_anBdh. i'' 1~~~~~~~~~~~~~~~~~~~~~~~ .Bgn47 '~~~~~~~~~~~~~~i > s _v.' 0 : s~~~~~~~~~ _~~~~~~~~~~~~~~~~~~~~~~~~~4 .AAM12/6 pe .~~~~~~~~~~~~~~~~~~~~~~~~ W.
not only by adding the abovementioned shrine to Cave 27 but by refurbishing the equally early Cave 25 and by starting the decoration of Cave 26's ambulatory. clearly part of the original patron's programmed work rather than something from the Period of Disruption. First of all. which had already been laid out a year earlier. work broke off very suddenly. for there was no apparent rush to complete it. Meanwhile. The latter is especially interesting. it was demonstrably the donation of Buddhabhadra: the plastering of the adjacent aisle ceiling. these Buddha groups include motifs that are characteristic of this very late level throughout the site. where he ordered the carving of the "gravastT Miracle" scenes. Equally significant. when the rest of the site was in such a frenzy of "finishing-up" activity. we can say that work ended on the cave at precisely that poignant moment when the Buddha was in the process of expiring. Cave 26 developed quite similarly. as was the similarly complex "late"image in Cave 26RW (fig. completed 478. . the connection of the Asmakas with the site-was cut off all of a sudden. in the oldand old-fashioned-Cave 27. incorporates previously cut areas into a new design. just as. During 478. another Sravasti Miracle scene at a point in the apse where the light was good. 13). like the now-defeated Upendragupta of Rishika. surely reflects the AMmakas'sudden assertion of independence. and the hugely moving Mahaparinirvaia. for the ceiling's plaster extends into the recess made by the panel. SA/29. SPINK V.Zza FIG26. However.eqa's death and which had been developing in a consistent way during that difficult year.Cave26 Left Wing. Begun 477. as indicated by the presence of unpainted sanguine sketches on the plastered background. had been one of Hariseqa's feudatories. And the evidence is not merely to be found in the dying Buddha. which had been spared the severe stresses that affected all of the other caves at the site during the year after Hari. plans were underway to create a new Buddha image in a newly added shrine (fig. patron of the cave. makes not even a ritual mention of the Vakataka overlordship. AAAUM 26LW Buddha (fig. obviously occurred after the Parinirvaxa scene was cut. not long after the weak new emperor had replaced his powerful father upon the throne. Only Buddhabhadra's donative inscription suggests an explanation. MainBuddha.34 Buddhabhadra's startling omission. s Se-ESO w. the Cave 26 inscription. the monk Buddhabhadra.32 The telling point is this: Buddhabhadra's own patronage-that is to say. which must have been proclaimed just after the various main Buddha images at Ajarta (including that of the Vakataka's chief minister) had been rapidly completed-that is. just as in Cave 16. seemingly. it too would surely have been completed. the formerly transcendent Guptas were put down by their increasingly independent feudatories. As we would expect. in this same era of decline. was continuing to develop his whole cave complex. Although the ruler of AMmaka. How do we explain this suddenly changed situation: that the Cave 26 complex. had there been but a few weeks' grace. 26).. That is. was brought to completion. the splendid Temptation by Mara. for the excavation of the shrine in the associated Cave 27 was cut off very suddenly too. this was never finished. so clearly an insult to the central imperial power. along with a handsome "Eight Buddha" sequence. the Parinirvar)a scene is unfinished. suddenly was afflicted too? Indeed. 27). work in the Cave 26 complex appears to have stopped even more abruptly than in any of those other caves where the Buddha images were being finished up during this anxious year. in the midst of work that could surely have been completed in a few days' time at most.33 The great house is denied its old honors completely. which. while celebrating the Mmaka rulers.90 WALTERM.
bythe end of 480. which Buddhabhadra's inscription so immediately reflects.16. But it does mean that the die had been cast. the hasty completion of the main Buddha images in Caves 2. anyone could put anything anywhere. Admittedly. That sad conclusion was yet to come. new shrine area (replacing cells). Starting in 479.aka emperor. the perniciously assertive AMmakas were ultimately responsible for the destruction of the great and ancient Vakataka house. Ajanta could by no means go on as before.Finaly. although in practice. the helter-skelter imagery of this period was added only to caves that had already been eA ~~~~~~~~~~~~~~~~x FIG. and the Ghajotkacha vihara. With things of the spirit now being urgently replaced by the needs of war. In such an environment. as well as the new emperor's weaknesses. and 22 atAjanta.21.4. AAAUM 49/78. aU such "disruptive' patronageends completely. and insurrection. for its prestigious patrons were the very people who would be most immediately affected by the developing crisis. Begtm 478. And the now even more suddenly suspended work in the more "privileged" Cave 26 complex and in the related "AMmaka" caves at Aurangabad records the crisis caused by the actual fruition of those aggressive intents-evidenced in Asmaka's sudden declaration of its independence. Cave 27. Instead. even on those of the locally ruling Mmakas. expansion. Intrusive images are now carvedor painted in all caves already dedicated. the sobering endings of the once exuberant programs of work at the Vakajaka sites do not mean that the central Vakajaka house had yet been overthrown. . it is no wonder that work on the Asmakas' huge cave complex stopped at the very moment that their less pious plans were made known. The next years were to be years of pain. the son of the great minister ofAsmaka-the very person in whose honor the Cave 26 complex was to be dedicated-had gone to the central Vakataka court to corrupt the heir apparent and insidiously prepare the way for the furdynasty's destruction. no further progress at all is made on the original programs of work in the caves.THE ARCHAEOLOGYOF AJANTA 91 The Dalakumaracarita tells us that once HariFera's successor was on the throne.27. for the whole of central India was ultimately caught up in the great insurrection that the Asmakas were fomenting. for the first time.36 Thus. immediately reflects the patrons' awareness of the Asmakas' aggressive intents. most of the other patrons had already rushed their Buddha images to completion and given up on the future some weeks or months before. U6.15. 7. Level5: Early479-Late 480 All consistentworkhas alreadystoppedon aU caves. by instigating a large coalition of the Vakataka's feudatories to rise up against the new Vakat.35 And the Dasakumaracarita ther states that. and one can only assume that in this atmosphere of aggression.
ende paroag . Significantly. the old Vakataka domain was in the hands of its former feudatories and others. No ^t supiigy rih n o rn aile 479utlzn eriercll deiaed(is 1920 w 2829)}o0r whee asin> Caes. who may well have got the diminishing group of painters or sculptors at the site to make these images for them in return either for prayers or for a pittance. Hariser)a's vast imperium. which by the mid470s had stretched from sea to sea. ". swallowed all that was theirs. often hastily inscribed with the brush.37 The Dasakumaracarita says that when the king of AAmakamaneuvered a great coalition of feudatories into rising up against the weak successor of the great Hariser)a and destroying him. . there was little hope that it would be restored in the foreseeable future. as we know from inscriptional evidence. SPINK A.sEIQ2 i'.Ntsrpiigyoeo theirst 28. And with the trade routes rapidly becoming unsafe for travel. Indeed. . Porch.But it was impossible for him to hold his greatprize together.Cave4. for a final moment. Whether or not this was true. AAAUM 861/68. "for the sake of spoils roused a quarrel among the chieftains and . the proud empire. left of doorway. we can certainly say that the economic base for the site had collapsed completely. including available locations inside the old Hinayina caves. Buddha (intrusive). The complete and sudden abandonment of the site's artistic patronage in 480witnessed by the host of half-completed contemporaneous images-stands as evidence of that expectation. what the foreseeable future offered as a prospect was only darkness. (fig 28. these (intrusive) images wpere onsrnet plce any patroailbesacsogheendedde (fig 28). Caves 26 and 19.fg_8.92 WALTERM.\ 1 L. the piirtiaghata of this great age in India. . which had so exuberantly and so briefly held aloft. was never to be heard from again. 1920 parng ende 2829 or whr (asiae som of. shattered into its componentparts.l ? N -~the first of -these intrusive images were placed on any spaces on thefcadeso available dedicte (figs. were made by the monks still resident at the site. but they rapidly occupied other spaces too. Late 479. the vast majority of these new dedications. surrisigly two chaiof halls. By the 480s and 490s. rear wall. he then. 29."9 FIG.
" a paper presented at The Age of the Vakatakas Seminar. We have again excluded the (presumed missing) image of Cave 8 from the discussion. 16. 26RW. For a general discussion of these developments see W. 1991). Yazdani. See W.. now published in R Parimoo et al. 12. For evidence that the donor's name was Upendragupta.1 (September 1990): 5-17 (containing an outdated time chart). The Buddha in the Ghatotkacha vihara could be counted as one more image that had to be quickly completed." Ars Orientalis 10 (1975): 143-69 (concerned with Cave 16). The inscription is replete with such 2. 6 for a brief discussion of the evidence. "Ajar)tA'sChronology: The Problem of Cave Eleven. The main image in Cave 26 and that in the right wing were also completed in normal course. and that he was of the local feudatory ruling house. 15. 1. 26LW. compare what remains on the porch pillars. but even if it hadbeen. 8." 143 n. Admittedly. Upendragupta surely would have dedicated his Cave 19 at the "eleventh hour" (actually at "one minute to midnight"). 17. and 29) discuss most of the other images under consideration here as well. 18. 'The Vakitaka's Flowering. originally following the pattern of those in Cave 20. see below. 10. however. 11. 19. 7. 1985. except in rather general terms. 26. 1. "The Vakataka's Flowering and Fall. were decorated while still in their blocked-out (square) format. see also n. Even the Buddha image in the leftwing may have been hurried to completion in the nervous climate of 478. Inscriptions of the Vdk7itakas(Ootacamund: Government Epigraphist for India. See n.19. in G. and later issues will contain my response to him and to A. 1981). See the monk Buddhabhadra's Cave 26 inscription.J. 1981). 1963). See n. see Spink. "Ajanta's Chronology: Cave 7's Twice-born Buddha. A "rebuttal" by Karl Khandalavala appears on pp. 26." Ars Orientalis7 (1968): 155-68. executed only after the doorway's carving had been completed in 468.ta (London: Oxford." in Kaladarsana. and was later much reworked. 4:114-18. since he could rather quickly 'complete" the cave in this way. "AjantA'sChronology: The Crucial Cave. Spink. 19. It is hard to believe that such an important record would have been merelypainted. the difference was hardly more than a matter of months. 3. V. Quoted from verse 24 of Upendragupta's Ajanta Cave 17 inscription. even if it was never inscribed. no traces of painted letters remain. Jamkhedkar. 5.21. but it appears that the patron did not continue much excavation at this point. and the fact that Cave 26 had been merely roughed out (rather than detailed) at this point.20." The seventeen are 1. with its 470-71-type projecting fitting. "Bagh: A Study. Spink. was left totally unfitted out. 14. 3. 11. In any case. Cave 8 must have had a loose image (now missing) because the stone at the lower level of the shrine is very bad. the whole rear area of Cave L6 was very rough when the crisis occurred. especially in the rear. The Cave 26 Left Wing Buddha image was probably made in the less up-to-date padmasana style because a serious flaw in the rock would have cut through the legs of a pralambapadasana image." Archives of Asian Art 30 (1976/77): 143-69. this probably occurred. The following articles by the author deal specifically with particular images discussed here: "Ajanta's Chronology: Cave I's Patronage. since Cave 26 was so intimately associated with the Asmakas. K Narain (New Delhi: Kanak Publications. 1955). L6. thiswould only confirm the need for haste. 2. so it may well have been used for anxious worship for a few days or weeks. TheArt ofAjanta (New Delhi: Vedam's Books International. A few cells were also unfinished at this time but were fitted out with later-style doorways after 475. 18-21 of the same number.3 (April 1991): 14-15. However. The painting of the left aisle wall was hurriedly started beforetheadjacent ceiling. . A. ed. 21. 22. 16. Mirashi. 109-26. This could explain why the shrine antechamber pillars. 4. 144-57. 26RW. Spink. "Ajantl's Chronology: Politics and Patronage. Spink. "Ajantl's Chronology: The Crucial Cave. Instead. he expediently plastered over these areas (possibly not even getting to paint the rear hall wall at this point). "Some Reflections on Professor Spink's Chronology at Ajanta. U6.THE ARCHAEOLOGYOF AJANTA 93 Notes This article is based on a paper presented at the Annual Conference of the Association for Asian Studies. Williams (New Delhi: Oxford and IBH in collaboration with American Institute of Indian Studies. are clearly unfinished. 7. ed. while the wall surfaces themselves are often abnormally rough beneath the masking plaster. when the cave became a focus of attention again in the Period of Disruption (479-80). 26LW. The five are 1. The small medallion on the beam connecting the two front center pillars does show distinct damage from use of its hook (now missing). The most thoroughjustification of this dating is in W. See the central porch medallion and doorway decoration. 1990. Various projecting portions at the lowerwall level. Chicago. Nagpur." ChhaviII (Banares: Bharat Kala Bhaven. being largely composed of friable red bole. it is reasonable to assume that work would have stopped on the Cave 26 complex not only very suddenly but prior to the time it stopped on all or most other caves. both having been begun before the wall (at its upper levels) was properly smoothed. 1985). 9. For discussion of the images of Cave 1 and 19.butboth had to be cramped into spaces different from those originally intended. 13. 2. 127-29. 103-16.4." Makarashtra Pathik 2. These and other articles (see nn. Even the decoration of the pillars is surprisingly hasty inside the cave. 15. makes it impossible to compare it to other caves as they appeared in 468. Ajan. An overview has most recently been published in Maharashtra Pathik 2. 6. only Cell R6. 7. 17. whereas the Buddhas in Caves 3 and 4A at Aurangabad would join the five that were finished in normal course. translated in V." in Studies in Buddhist Art of South Asia. Actually.
for the majority of the panels are intrusive. In some other cells. forthcoming 1992. Dharmadatta. curiously but perhaps suggestively. it is quite possible that it had not been completed when the Asmakas' control of the cave ended and that the doorway was therefore finished in the more uncontrolled clime of 479. 'Politics and Patronage. Even if this is so. Ryder. were responsible for Harisera's death. 1962).the son of the Aimaka minister came to the Vakataka court 'pretending to have been exiled by his father. werejust sliced away to the degree necessary to allow the new recessed mode to be cut. but the very lack of wear and tear and 27. Satakopan et al. generally proceeding from front to back. 19. 152. 1967). Intrusions were never placed in or on the caves until the main image had been dedicated and put into use. Dan4in's Daiakumaracarita (Madras:V. The projections in Cave I's rear cells. One possible exception is on the shrine doorway of Cave 26LW. verse 10. C. but we do not. This is most evidentfrom the fact that the "normal" progress of work continued on them during 478. suggests that they began to be used as soon as they were habitable-even before the cave was dedicated. suggest that these caves were the first two put into ritual use. 1. 30. 23. Majumdar and A. C. This assumes that the group that was plugged in was as conventional in size as in placement. The heavy deposits of grime in Cave 17. as can be seen by studying them in connection with table 2. which had not even been supplied with their pivot holes when the Hiatus occurred. in the intrusive Buddha images placed in Cave 16 by Buddhabhadra's associate.. 100). and 24. "who accomplishes. as we can tell by the wear in various cell doorway pivots. although the date of this is uncertain. A mere "census" of the site makes this clear. See n. Indeed." 26. This byno means suggests that the Ahmakas. Had controls been relaxed during the Hiatus. see Spink.. Monksobviouslycontinued to live in the cavesfor at least a decade or so after 480. See the Daiakumaracarita." as well as to his generosity. "Ajarqa's Chronology. obviously Varlhadeva no longer controlled the cave by this time. 21. Perhapsthe caveswere finally abandoned when the AAmakas. where one of the multiple Buddhas framing the doorway has his proper right hand in abhaya mudra. Since there are traces of color elsewhere on the Pavinirv&aa scene. 24. 7. W. scarcely a year later. 37. referred to in Buddhabhadra's inscription (n. 28. 8). 1 for discussion. S. even such tasks as would normally call for rigours and active struggle. Admittedly.) 36. with tact and sweetness only. See R. Insc7iptions. 1927]. despite their plottings. The V&k&*takaGupta Age (Delhi: Motil al Banarsidass. In both Cave 7 and Cave 21. as could have been the case if the image here had been started by the original donor in 478. it is conceivable that this area was painted too but that the surface paint has fallen away. sometimes placed in the courtyards of unfinished caves or in unused areas of scarp between the caves. but on the other hand a poisoning could easily have gone unrecognized. the old projections were abandoned but left intact when the doors were refitted in the new mode after 475. The father of this "son of the minister of Asmaka" must have been the current minister. all obvious intrusions at the site "fit"into a much later period (479-80). RamaswamySastrulu and Sons. This conclusion accords with our "rule" that intrusions (of which this cave has many) were placed only in caves where the image had already been dedicated. it is conceivable that new donors took over the abandoned cave in 479 and treated it as if it were their own shrineletas they did in the court "shrinelets" (originally cells or cell complexes) of Caves U6. According to the Da'sacuWaTracaGita. the preparatory plastering for the paintings done at this time was applied over surfaces that were not yet fully smoothed. A. 31. See n. Spink." fig. like those in Cave L6. 22. 20. See n. See my article. or was about to be dedicated. 32. the ambulatory decoration was evidently abandoned in mid course when time ran out in 478. 145. lost control of the region. 39. that 'he filled the life of a man with worthy deeds. "ARecently Discovered Buddha Image at AurangabAd. Altekar." for the R. at the behest of the monks still remaining at the site. 38. 25. the manner in which the cells were sequentially fitted out. The Dalakum&raca7itasays. Mirashi. ed. This explains how he could have been accepted there even after AAmaka had renounced its ties to the imperial house (in 478). 189ff. In Cave 17. Agrawala grime in such caves suggests that this period of residence was verylimited." He seems to have bequeathed some of these qualities of deviousness and subtlety to this son. SPINK references to Upendragupta's trust in 'the power of the expenditure of wealth. in 478). 8. for an example." See V. (Images were. we would surely find intrusive images interrupting the programmed work done after 475. clearly because of a flaw where his hand would otherwise be.e. by virtue of which he "caused thejoy of suppliants by conferring gifts (on them). yet for the unworthiness of his subjects was translated into divinity" (trans.94 WALTERM." Felicitation Volume. 34. . The AAmakas' rout of the Vakitakas is further reflected. 39. TheTenPrinces [Chicago: University of Chicago Press. in 471 and 468 respectively. with their Buddhist inclinations. 33. but the iconographic and stylistic connections between them and Cave 26 are also striking. however.. 35. Since this doorway (like that of the porch) was being decorated verylate (i.
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Sequencing genes in silico using single nucleotide polymorphisms
The advent of high throughput sequencing technology has enabled the 1000 Genomes Project Pilot 3 to generate complete sequence data for more than 906 genes and 8,140 exons representing 697 subjects. The 1000 Genomes database provides a critical opportunity for further interpreting disease associations with single nucleotide polymorphisms (SNPs) discovered from genetic association studies. Currently, direct sequencing of candidate genes or regions on a large number of subjects remains both cost- and time-prohibitive.
To accelerate the translation from discovery to functional studies, we propose an
Prior to the general availability of routine sequencing of all subjects, the ISS method proposed here provides a time- and cost-effective approach to broadening the characterization of disease associated SNPs and regions, and facilitating the prioritization of candidate genes for more detailed functional and mechanistic studies.
The recent advent of SNP array based high throughput genotyping technologies has stimulated a wave of genetic association studies, including genome-wide association studies (GWAS)
In this study, we describe an in silico sequencing (ISS) method to more fully define the sequence variations in candidate gene regions. Recently, the 1000 Genomes Project
This development is an extension to our recently developed method
This method of predicting polymorphic genetic variants is closely related to the commonly used imputation methods. Typically, imputation methodologies such as Impute
In the remainder of the paper, we first describe the proposed methods then summarize the properties of genes deep sequenced in the 1000 Genomes Project, and evaluate the performance of our gene prediction models in the Results Section. Then we present an application of our approach to an existing GWAS data gathered from the WTCCC. We conclude this paper with discussions on the strengths and limitations of the proposed method.
Deducing Diploid Sequences
In order to build prediction models for gene diploid sequences, one of the first steps is to deduce the phase information of the gene sequences for the training dataset, which are the samples sequenced in Pilot 3 of the 1000 Genomes Project here. Utilizing all SNP calls within each gene, we deduced diploid sequences for every subject. This is equivalent to phasing multiple SNPs, in the absence of any structural variants as in the current case. Several methods have been developed for phasing multiple SNPs
Predicting Multi-Allelic Genes
With these phased gene diploid sequences, we expanded the previously developed methodology
which leads to
under the Hardy-Weinberg equilibrium (HWE). The detailed estimation procedure was described in
We build a procedure to systematically select the most informative SNPs using a forward-and-backward scheme, which starts with the SNPs within a gene (if available) and gradually extends to flanking regions of each gene. The SNP selection process was evaluated by an objective function, which is the negative log-likelihood of the gene allele given SNP genotypes penalized by the number of additional parameters to be estimated (Akaike information criterion (AIC)
where the second term equals the difference of the number of gene-SNP haplotypes (m) and the number of gene alleles (k).
The search region was empirically chosen to maximize the prediction accuracies while retaining parsimony. We have chosen 150 kb flanking region to predict HLA genes in our previous work
Additional file 1
Supplement Methods. SNP selection process in building prediction models.
Click here for file
Prediction accuracies
As part of the model validation, we applied prediction models built from the training set, to predict diploid sequences on another dataset, and to compare the predictions with the actual diploid sequences. For this comparison, we used the following accuracy measurement. Suppose that
represents the number of correctly predicted diploid sequences for the ith subject.
Measuring genetic polymorphism
Typical genes under consideration are polymorphic, i.e., with multiple alleles. To measure such polymorphisms, we propose to use the Shannon entropy, which is commonly used to measure variability of discrete variable
where the summation is over all possible alleles. Entropy with value of 0 means that the gene is not polymorphic. The maximum value of entropy equals to
Analysis Pipeline
Here we describe a general analysis strategy for building prediction models for diploid sequences of genes sequenced in the 1000 Genomes Project using SNPs. The resulting prediction models can be easily applied to sequence candidate genes in silico and facilitate the interpretation of association discoveries. To build reliable prediction models based on a relatively small number of samples from each population (Table
Table 1
Samples genotyped in the 1000 Genomes Project pilot3 and the HapMap3 Project
1000 Genomes
Both projects
Project pilot3#
East Asian
West African
# The 1000 Genomes project pilot3 sequenced two trio samples from CEU and YRI. The children in these families were excluded from this analysis.
* Only samples that were in the same 7 populations of the 1000 Genomes project were listed here. The HapMap3 project also genotyped samples from ASW (African ancestry in Southwest USA), GIH (Gujarati Indians in Houston, Texas), MEX (Mexican ancestry in Los Angeles, California), and MKK (Maasai in Kinyawa, Kenya).
Given the large amount of sequencing data, the typical first step is to align short-read sequences to the reference genome, and then to make variant callings. In this project, investigators from the 1000 Genomes Project have already performed such analysis, producing all possible SNP genotype calls in each study population.
The second step is to clean the data. For each gene, we filtered out samples that have at least one SNP missing in the gene. Genes that have more than 50% of samples filtered or have only one SNP were excluded from analysis. In addition, samples that were filtered in more than 60% of genes were excluded.
The third step is to deduce diploid sequences by phasing genotypes of all SNPs within every gene for every subject. To maintain phasing efficiency and at the same time retain the independence of the training and validation sets, we estimated the frequency of alleles of each gene using the entire set of samples, and then predicted the phases of gene alleles for samples in the training and validation set separately. Each distinct gene allele was then assigned a label as GeneName*Allele, where Allele was labeled by the numerical number 1, 2,..., in the order of the frequency of corresponding gene allele. The diploid sequences were called for each subject, if the corresponding posterior probability was maximum and also exceeded a pre-assigned threshold. Otherwise, the subject's sequence pair was deemed as missing. To ensure the quality of prediction models, a stringent threshold of probability > 0.95 was applied and only genes that had less than 10% samples with missing diploid sequences in both training and validation sets were selected for further model building.
The fourth step is to build prediction models for each gene using our algorithm of predicting multi-allelic genes. The prediction models built based on diploid sequences and SNP genotype data of samples in the training set were then applied to the training set itself. The prediction accuracy on the training set was thus evaluated by comparing the predicted diploid sequences with the observed ones. To control the quality of prediction accuracy, we used call threshold (CT) of CT = 0, 0.5 and 0.9, i.e., one would make a prediction only if the posterior probability exceeds CT.
The last step is to validate these prediction models. By applying the prediction models built in the above step to the validation set, we computed the prediction accuracy using CT = 0, 0.5 and 0.9, which would reflect "true" prediction accuracies in practice.
Deeply Sequenced Genes
All variants from Pilot 3 of the 1000 Genomes Project have been identified and organized in a variant file
Figure 1
Categories of genes targeted in the 1000 Genomes Project Pilot3
Categories of genes targeted in the 1000 Genomes Project Pilot3. Of the 1,000 CCDS genes targeted, 93 genes (Missed) were not targeted successfully in at least one genotyping center, 45 genes (0 or 1 SNP) have only zero to 1 SNP called, 219 genes (Filtered) were filtered out (≥ 50% of the samples have missing value in at least one SNP within the gene), 32 genes (Unphased) were not phased properly (≥ 10% of the samples have phasing probability < 0.95 in either training or validation set), the remaining 611 genes (Predicted) were selected for analysis in this project.
Properties of Selected Genes
Utilizing all SNPs in the gene, we successfully deduced diploid sequences for 611 of the 643 genes (See Methods Section for further information). Of these genes selected for the following modeling exercise, 84% are less than 100 KB, while the remaining are larger, with the largest being 856 KB in size (Figure
Figure 2
Histograms of the characteristics of genes (N = 611) selected in this project
Histograms of the characteristics of genes (N = 611) selected in this project. Note that the interval length of each group is not evenly distributed.
Additional file 2
Supplement Table 1. Genes and their Properties.
Click here for file
Training Prediction models
Of the individuals sequenced in Pilot 3, 614 samples from multiple populations have been genotyped in HapMap 3 using the Affymetrix Human SNP array 6.0 and the Illumina Human 1M-single BeadChip. We have recently shown that prediction models built using a training set that includes multi-ethnic populations are reliable and perform well for the subpopulations
Using the training set, we built prediction models for all 611 genes and plotted the rescaled objective function value Ôg (See Methods Section for further details) for each gene at each flanking region size, quantified by the color intensity as shown in the color bar. As illustrated in Figure
Figure 3
Plot of rescaled objective function value versus the flanking region size
Plot of rescaled objective function value versus the flanking region size. The prediction models for the set of 611 genes selected were built using nine different flanking region sizes, shown on x axis. The y axis shows the list of genes (name not shown). For each gene g, the objective function value
Upon the completion of the model building, we predicted diploid sequences for the training set using prediction models, and compared prediction results with observed diploid sequences. Using call threshold (CT) as 0, 594 out of 611 genes have predictive accuracies averaging 98.26% (range, 95% to 100%) (Figure
Figure 4
Prediction accuracy (%) and Call rate (%) of prediction models built using flanking region size 250 K in the training set (N = 342) and the validation set (N = 342)
Prediction accuracy (%) and Call rate (%) of prediction models built using flanking region size 250 K in the training set (N = 342) and the validation set (N = 342).
Prediction accuracies were examined in the relation to entropy (Figure
Figure 5
Relationship of the accuracy of prediction models with gene entropy in the training set and the validation set at CT = 0
Relationship of the accuracy of prediction models with gene entropy in the training set and the validation set at CT = 0.
Validating Prediction models
The prediction models built from the training set were validated by predicting sequence variants for each of the 611 genes in the independent validation sample set. The accuracy of the prediction was evaluated by comparing between the predicted and observed diploid sequences. The distributions of prediction accuracy with CT = 0, 0.5 and 0.9 are illustrated in Figure
Additional file 3
Supplement Table 2. Accuracy % (call rate %) of the prediction models evaluated on training and validation set.
Click here for file
Final Model
The training and validation exercises described above have shown that the proposed method of in silico sequencing candidate genes is valid, scalable, and has satisfactory prediction accuracies for a broad sampling of genes. To make the gene prediction models more robust, we combined the training and validation sets to build the final prediction models for all 611 genes using the procedure described above. The prediction models included the number of SNPs ranged from 1 to 38. Those models and the SNPs selected are available on our website
KCNJ11: A Type 2 Diabetes Associated Gene
To illustrate the utility of the prediction model in silico gene sequencing, we analyzed the Type 2 diabetes (T2D) GWAS data generated by WTCCC
The predictive model was built for KCNJ11 gene following the above modeling process, using the available SNPs from HapMap 3. Since not all selected SNPs were genotyped in WTCCC, we employed IMPUTE v2 to impute the additional SNPs, using default parameters. With both the imputed and genotyped SNPs from WTCCC, we sequenced the candidate gene KCNJ11 in silico based upon the predictive model. The distribution of predictive probabilities is illustrated Figure
Figure 6
Distribution of predictive probabilities (the highest posterior probability among predicted KCNJ11 alleles for every subject) in the WTCCC data
Distribution of predictive probabilities (the highest posterior probability among predicted KCNJ11 alleles for every subject) in the WTCCC data.
Nine gene alleles were predicted to be in both case and control sets. Using the most frequent allele (KCNJ11*3) as the reference, we performed a multi-allelic association test using HPlus
Table 2
Results of the multi-allelic association of the KCNJ11 gene alleles with T2D.
(0.8, 0.98)
(0.8, 1.52)
In WTCCC1 cohort, which have 1924 T2D cases and 2938 controls (NBS and 58BC), a total of 9 gene alleles were predicted with variations at 9 SNP locations (rs41282930, rs8175351, rs5215, rs1800854, rs1800467, rs5216, rs5218, rs0#, rs5219). The 4 common gene alleles (freq > 0.01) were analyzed in a multi-allelic association in this case-control study.
*The SNP variations among the common haplotypes (freq > 0.01) are highlighted in bold.
#This SNP is located at chr11:17366025 (NCBI build 36) and has not been assigned an rs number.
To investigate the functional basis of these variants, we examined the amino acid sequence variants for KCNJ11 gene. Utilizing the reference genome from NCBI
Table 3
Results of the multi-allelic association of the KCNJ11 amino acid gene alleles with T2D.
(0.9, 1.69)
By converting the predicted genomic gene allele to amino acid sequences, the total number of gene alleles is reduced to 5, with four amino acid variations (E23K, L64V, L270V, I337V, and S385C). The 3 common haplotypes were considered in this case-control study.
*The amino acid variations among the three common haplotypes are highlighted in bold.
The 1000 Genomes Project, built upon the success of the HapMap project, promises to generate high quality sequence data for thousands of subjects from multiple racial/ethnic groups. Besides shedding light on human genome variations, this project will also generate an abundance of sequence data that will be useful for fine mapping and better defining disease associated functional variants. Towards this goal, we have described a method for in silico sequencing of candidate genes and regions of potential interest. Through training and validation exercises on currently available sequence data from the 1000 Genomes project, we have shown that the method produces reliable prediction models with high accuracies, with few exceptions. To facilitate the future use for the research community, we developed a set of predictive models for 611 genes utilizing all available sequence data. Such models can now be used to sequence known candidate genes in silico and further to investigate their functional properties, prior to carrying out actual sequencing in the wet lab. Therefore, applications of this method would help accelerate validation researches from discoveries to functional characterization in the most cost-effective and time-efficient way.
Connection with SNP Imputation Methods
Our method of predicting polymorphic genetic variants may be thought of as an extension to imputation methods. Specifically, like imputation methods, our method also uses local LD to infer untyped gene alleles (or equivalently, phased diplotypes with multiple SNPs), whereas imputation methods infer untyped SNPs. By predicting fully phased sequence diploids, our approach allows one to translate DNA sequence variants in the coding region into amino acid sequence variants, thereby facilitating the functional interpretation of disease associations.
Connection with Haplotype based Association Analysis
Our methodology also helps to resolve a long-standing debate on the value of haplotype based association analysis. Recall that haplotype based association analysis, since its inception, has been suggested as a preferred method for assessing disease association
Structural Variants
The current methodology can naturally incorporate structural variants in building prediction models. One of the major advantages of direct sequencing is the ability to identify structural variants, including insertions and deletions. Structural variants could be naturally coded as additional gene alleles, and incorporated into prediction models. In the current study, few genes are known to have structural variants in those sequenced exons; no illustrations are given here.
Choice of the Flanking Regions
During the model training process, one subjective parameter to be chosen, is the flanking region size for each gene (or region). The larger the flanking region, the more likely it is that all predictive SNPs are included, but also the greater is the chance that the prediction models will be over-fitting. Based upon our empirical experiences determining the optimal size of flanking regions for predicting alleles for five HLA genes
Limitation 1: poor prediction accuracies
While prediction accuracies of most genes are approaching 99%, it is noted that two genes (TLL1 and ZNF474) have particularly low prediction accuracies in the validation set. At CT = 0, the lowest prediction accuracy among all the 611 genes is 79% for TLL1 gene. This gene is one of the complex genes in this dataset (see Figure
The second gene ZNF474 has the prediction accuracy of 80% on the training set. Seventy out of the 92 prediction errors involved three alleles ZNF474*1, ZNF474*2 and ZNF474*3, which differ by two SNPs (rs2560306 and rs35262183). Unfortunately, neither SNP was genotyped nor in significant LD with any of the SNPs in the HapMap 3. In the absence of these "highly informative and yet isolated SNPs", one would not be able to differentiate these alleles. This phenomenon has also been observed for HLA predictions
Limitation 2: poor diploid sequence quality
In order to recover diploid sequences from the 1000 Genomes Project, it is necessary to statistically infer for haplotypes. Among the 645 selected genes, 32 genes have been excluded, because > 10% of samples had relatively poor haplotype inference with probability less than 0.95. The poorer haplotype inference may associate with a complex gene structure. For example, of these 32 genes, four genes (ANKRD15, TRPV3, NLRP11, CYP24A1) are longer than 850Kb, or the entropies are greater than 2, or have more than 50 gene alleles. To address this challenge, our strategy is to divide a gene into several regions, e.g., by combining adjacent exons or by short segments, and then deduce the alleles within each region separately. On the other hand, for those genes with complex structural variations, probably the most effective approach is to use sequence technologies that produce relatively long reads, such as several hundred or even thousand nucleotide bases, which allow us to deduce diploid sequences directly. Technological improvement in the future would lessen the impact of this particular limitation.
Limitation 3: limited polymorphisms in the training samples
A critical factor of building useful prediction models, especially for those genes with rare gene alleles, is to have a sufficiently large sample size so that rare alleles are observed in the training set. In our study, the SNP and sequence data used in this paper came from the 614 samples in both HapMap and the 1000 Genomes Project. Prediction accuracies should be improved when much more samples are available in future.
Prior to the general availability of routine sequencing all subjects, the ISS method proposed here provides a timely and cost-effective approach to broadening the characterization of disease associated SNPs and regions, and facilitating the prioritization of candidate genes for more detailed functional and mechanistic studies.
Authors' contributions
X.C.Z. participated in the initial discussion of this project, performed all data analysis, and participated in drafting the manuscript; B.Z. contributed to the development of ISS software. S.S.L. contributed to assisting the data analysis, and to preparation of the manuscript; X.H. contributed to the discussion of methodology; J.A.H. contributed to the original idea, and helped the manuscript drafting; L.P.Z. conceived the idea, and lead the project. All authors read and approved the final manuscript.
Acknowledgements and funding
We would like to thank NIH for providing partial funding support to this project through the following grants: NIH/NIMH R01MH084621 (PI: Lue Ping Zhao), NIH/NCI R01CA106320 (PI: Lue Ping Zhao), NIH/NCI R01CA119225 (PI: Lue Ping Zhao), and NIH/NHLBI R01HL087690 (PI: John Hansen).
Genomewide Association Studies and Assessment of the Risk of Disease
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subset: commoncrawl