ࡱ>  0bjbj AN'a %%%8]T$%w*t)))))))$+.*!!!*X1*(((!)(!)(((@%$N()G*0w*(9/&9/((9/(`[( \iS**'w*!!!!9/ : Barley Genetics Newsletter (2009) 39:20-23 Historic barley seed samples and their utility M. W. Leino1* and J. Hagenblad2,3 1 Swedish Museum of Cultural History, SE-643 98 Julita, Sweden 2 Uppsala University, Department of Ecology and Evolution, SE-752 36 Uppsala, Sweden 3 Norwegian Univ. of Science and Technology, Dep of Biology, N-7491 Trondheim, Norway *Author for correspondence: Matti.Leino@nordiskamuseet.se Gene banks around the world hold over 370.000 accessions of barley (van Hintum and Menting, 2003). Although a large number of landraces and obsolete cultivars are stored in genebanks the geographic coverage of this conserved material is far from evenly distributed. For example only a handful of landraces from Northern Europe have been conserved (Jones et al, 2008a). Thus studies of geographic differences in land race barley are hampered by the availability of suitable material. Another major drawback with gene bank material is the authenticity of the available accessions. Evolutionary forces such as drift and selection may have affected the genetic diversity since the accessions were deposited in gene banks and there is also a risk of contamination from poor propagation practices (Parzies et al., 2000). Furthermore, the true geographic origin of cultivars may be unknown or uncertain. Recently material from non-viable historic collections has become recognized as an alternative to the use of gene bank material. Herbarium material can be well documented and therefore associated with useful geographic information, but the amount of material available is often limited and successful genotyping can prove problematic (Savolainen et al., 1995). In contrast, seed collections can contain equally well-documented samples with large number of seeds from each accession, thus enabling studies on a population level. Moreover, DNA quality of aged seeds widely surpasses that of other equally old vegetative tissue like straw, leaves and chaff (Lister et al., 2008). In Sweden there are two large seed collections, both assembled at the turn of the last century. The collection at the Swedish Museum of Cultural History contains 419 different accessions of barley from 1865 1918 (Leino et al., 2009). These samples were collected on farm from many different counties across Sweden and Finland with a minor addition of accessions from the rest of Europe (Fig. 1). Another seed collection is stored at Vnersborg Museum (Johansson, 2003) and contains 61 barley accessions. The samples in this collection originate from an agricultural exhibition in 1882. The seeds are in both cases stored in sealed glass containers and were displayed on exhibitions and museums but have never regenerated (Fig. 2). We have genetically analysed part of the collection at the Swedish Museum of Cultural History (Leino et al., 2009). The DNA of these 90 to 140-year-old seeds is severely degraded, but in spite of this degradation PCR amplification is possible, in particular of fragments less than 300 bp long (Fig. 3). We have successfully amplified not only multi-copy genes such ribosomal DNA, but also single copy genes like microsatellites.  Figure 1. Barley harvest in the county ngermanland, Sweden in 1905. Photo: Petrus Hedstrm, Swedish Museum of Cultural History.  Figure 2. Examples of glass containers with barley seed stored at the museum.  Figure 3. Analysis of DNA quality in fresh and historic barley seed. DNA isolations are made from single seeds. A) Electrophoresed total DNA. B) PCR amplified microsatellite marker fragments. M=DNA size marker, F=fresh seed, H=historic seed. Recently, we have conducted a population genetic study of Swedish landrace barley using 113 year-old seed samples (Leino and Hagenblad, 2009). Our aim was to investigate the effects of bottlenecks, adaptation and gene flow on genetic diversity within and between landrace populations. The results demonstrated varying levels of within-population variation with some decline in diversity for more Northern populations. We could also detect clear population differentiation, and population structure within Sweden into a Southern and a Northern cluster. These results possibly reflect different introduction routes of barley into Sweden. Historic material has also been used to study the geographic distribution of functional genes. In a study of the flowering time gene Ppd-H1 in gene bank material originating across Europe, Jones et al. (2008b) showed geographic differentiation, with one allele gradually replacing the other when going north across Europe. Lister et al. (2009) repeated the study using historic samples, including several samples from the Vnersborg collection. In this study a much clearer picture was reported with one allele being present north of the Alps and the other in southern Europe. We have found the same pattern in this gene with both alleles being present in gene bank material originating from Sweden, but only a single allele present in historic material from the same region (Knppel, unpublished). This further illustrates the presence of ambiguous material in genebank collections. Non-viable historic barley material is a valuable complement to extant gene bank accessions, especially for geographical areas where gene bank material is rare or non-existent. Further studies of neutral and functional markers may shed additional light on the history of barley cultivation and breeding. It is thus imperative that historic material receives the same conservation attention as extant material in our gene banks. References: Hintum, Th. van, and Menting, F. 2003. Diversity in ex situ genebank collections of barley. In: Bothmer, R. von, Hintum, Th. van, Knpffer, H., Sato, K. (eds) Diversity in Barley (Hordeum vulgare). Elsevier Science B.V., Amsterdam, The Netherlands. p. 29-52. Johansson, P. 2003. Prof av vrt svenska fr- Vestra Sveriges landtbruks-museum. In Johansson, P., Aarsrud, S., berg, A.-C. (eds.) Det underbara huset: Vnersborgs museum. Vnersborgs museums skriftserie 1:115-122. Jones, H., Lister, D.L., Bower, M.A., Leigh, F.J., Smith, L.M.J., Jones, M.K. 2008a. Approaches and constraints of using existing landraces and extant plant material to understand agricultural spread in prehistory. Plant. Gen. Res. 6:98-112. HYPERLINK "http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Jones%20H%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus"Jones, H., HYPERLINK "http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Leigh%20FJ%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus"Leigh, F.J., HYPERLINK "http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Mackay%20I%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus"Mackay, I., HYPERLINK "http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Bower%20MA%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus"Bower, M.A., HYPERLINK "http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Smith%20LM%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus"Smith, L.M., HYPERLINK "http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Charles%20MP%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus"Charles, M.P., HYPERLINK "http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Jones%20G%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus"Jones, G., HYPERLINK "http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Jones%20MK%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus"Jones, M.K., HYPERLINK "http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Brown%20TA%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus"Brown, T.A., HYPERLINK "http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Powell%20W%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus"Powell, W. 2008b. Population-based resequencing reveals that the flowering time adaptation of cultivated barley originated east of the Fertile Crescent. Mol. Biol. Evol. 25:2211-2219. Leino, M.W. and Hagenblad J. 2009. 19th century seeds reveal the population genetics of landrace barley (Hordeum vulgare). Mol. Biol. Evol. in press. Leino, M.W., Hagenblad, J., Edqvist, J., Karlsson Strese E.-M. 2009. DNA preservation and utility of a historic seed collection. Seed Sci. Res. 19:125-135. Lister, D.L., Bower, M.A., Howe, C.J., Jones, M.K. 2008. Extraction and amplification of nuclear DNA from herbarium specimens of emmer wheat: a method for assessing DNA preservation by maximum amplicon length recovery. Taxon 57:254-258. Lister, D.L., Thaw, S., Bower, M.A., Jones, H., Charles, M.P., Jones, G., Smith, L.M.J., Howe, C.J., Brown, T.A., Jones, M.K. 2009. Latitudinal variation in a photoperiod response gene in European barley: insight into the dynamics of agricultural spread from historic specimens. J Arch. Sci. 36:10921098. Parzies, H.K., Spoor, W., Ennos, R.A. 2000. Genetic diversity of barley landrace accessions (Hordeum vulgare ssp. vulgare) conserved for different lengths of time in ex situ gene banks. Heredity 84:476-486. Savolainen, V., Cunoud, P., Spichiger, R., Martinez, M.D., Crvecoeur, M., Manen, J.-F. 1995. The use of herbarium specimens in DNA phylogenetics: evaluation and improvement. Pl. Syst. Evol. 197:87-98.     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Hagenblad,22(H,1,22,2% % % T` Dc @E@ LahT2,3"!% % % TTd D @E@d LahP  % % % TT  @E@ LahP  % % % TX*u@E@fLahP1 "% % % T0:? @E@&LahSwedish Museum of Cultural History, SE8H,2'2Y2',2N2!C12!,H'2"08=TT@ :` @E@@ LahP-!Ta :@E@a Lahx643 98 Julita, Sweden22223(2,8H,2,2TT:@E@LahP - % % % TX@E@LahP2 "% % % T @E@;LahUppsala University, Department of Ecology and Evolution, SEH22',,H22,!'0I-2,!N,22!=,2330,22=222228=TT  @E@ LahP-!T @E@ Lahx752 36 Uppsala, Sweden22222H22',,8H,2,3TT@E@LahP - % % % TXg[@E@gLLahP3 "% % % T S@E@z Lah`NorwegianH2!H,1,2TTT m@E@TzLahP Tln N@E@nzLahXUniv.H22TTO g@E@OzL&" WMFC "2222,NH2!H.1TT %@E@zLahP -!ah'% % "  T@E@9Lah*Author for correspondence: Matti.Leino@nordiskamuseet.se2H222!!2!,3!!,'2223-2,,Y,<,22\22!2(2,N2',,'-TT@E@LahP - TTO{w@E@O`LahP - TTOy{@E@OLahP - TTO{]@E@OFLahP - TdO_ @E@OLahTGeneH,2,TT!_Q@E@!LahP 1TR_\@E@RLahbanks around the world hold 3,22'1,!222213-1H2!212221T]_ @E@] Lahdover 370.00022,!1222322TT _ @E@ LahP 1T _]@E@ Lahxaccessions of barley (,,-,''22'12!12,!.11!T^_+@E@^Lahlvan Hintum and 2,21H22N1,22 TOC @E@O, LahhMenting, 2003Y,221'2222T0 C @E@, &Lah). Although a large number of landrace!'H22212',',"2,'22N2,!'2!'-22!,,,T8 *C @E@ , 'Lahs and obsolete cultivars are stored in '',22'22'2,,',22,!'',!,''2!,2'2 T|OE + @E@O ]Lahgenebanks the geographic coverage of this conserved material is far from evenly distributed. 1,3,2,22(2, 2,31!,22,,23,!-2,3!2',22(,"2,2N,-!,'","!!2O ,2,202'!22,2 T\O ) @E@O XLahFor example only a handful of landraces from Northern Europe have been conserved (Jones 72!,3,N2,22/,2,23!22!,22!,,,'!!2NH3!2,!2=2!22,2,3,2,,2,22',!2,2!(22,'T` *) @E@ LahTet , T`O+  @E@O LahTal,,TT+  @E@ LahP &Td+  @E@ LahT20082222TT+  @E@ LahPa,T+  @E@ Lah|). Thus studies of geogr!&=22'&'22,&&2!&1,31!T+  @E@ Lahxaphic differences in ,22,&2!!,",2-,'&2&T + @E@ Lah`land race ,22&!,,,&T + + @E@ Lahbarley are hampered by the 2,!./',",&2,N2,!,2&4/&2- TO F @E@O "Lahavailability of suitable material.,2,,202!'2,2,O,,!,TTG s @E@G LahP - TTO { @E@Ok LahP - T O ` @E@O LahAnother major drawback with geneH222,!;N,2";2",H2--2;H2<1,3,TTa  @E@a LahP ;T + @E@ 3Lahbank material is the authenticity of the available 2,&" WMFC 2,',T| 'X @E@ Lah\samples ',N2,'TY '@&" WMFC <<E@Y Lahlwere collected I,!,,2,-,2T'O@E@ Lah` on farm !22!-!N!TlP'+@E@PLahXfrom !!2N TO @E@ODLahmany different counties across Sweden and Finland with a minor additN,302!",!,2,222,',,!2''8H,2,2,2272-22H2,N22!,22Tx  @E@ Lah\ion of 212!T1 @E@ Lah`accessions,,,,''22'TT2N @E@2LahP TlO+ @E@OLahXfrom !!2ORp @Times New Romanԓ ғ P`2ԓ ѓ ӓ ԓ O`2ԓ ѓ jEU1ѓ ԓ ܳCFU1XG* Times ew omanѓ PM1ғ ғ =RK1,ғ ܳCdvdv% Rp@Times New RomanܳCFU1Xϓ |[Г A|h]||ϓ  @w=!%d Z(XX   P Г .["tГ |[\ѓ ѓ wYwdv% ( % (  TO ~@E@Og Lahhthe rest of E2,9!,'92!9=T N~@E@gLahlurope (Fig. 1). 2!23,9"7192!:T8O ~@E@Og'LahAnother seed collection is stored at VH222,!9'-,29,3,,229'9'2!,29,9H,T *~@E@gLahlnersborg Museum 2,!'22"19Y2(,2O TO @E@O Lah`(Johansson!(22,2''22Tx *@E@ Lah\, 2003)2222!TT+G@E@+LahP TH`@E@H Lahhand contains ,22-22,2'T`a@E@aLahT61 22T @E@Lahpbarley accessions2,!.0-,,,''22'TP +@E@ +Lah. The samples in this collection originate =2,',N2,(22',2,,222!12,, TDOd@E@OM)Lahfrom an agricultural exhibition in 1882. !!2N',2(-1!,22!,(,32222'2'2222'TN d@E@MLahhThe seeds are =2,'',-2'',!,'T,O +d@E@O M%Lahin both cases stored in sealed glass 3'222',,(,'''2!,2'2''-,,2(1,'( TDOfQ@E@OTLahcontainers and were displayed on exhibitions and museums but have never regenerated ,22,2,!' ,22 I,", 2'2-0-2 22 ,32222' ,21 N2',2N' 22 2,2, 2,2-! !-1,3,!,,2 TTRfr@E@RLahP(!Tlsf+@E@sLahXFig. 71 TTOJ@E@O3LahP22TXJ@E@3LahP).!TTJ@E@3LahP - TTOL{@E@OLahP - TPO+0@E@OVLahWe have genetically analysed part of the collection at the Swedish Museum of Cultural _,/2,2,01,3,,,0/,3,0',3/2,!/3!/2,0,2-,22/,/2,/8H,2'2/Z2',2N/2!/C22!,Rp@Times New Roman 8ѓ ԓ ғ P`2ԓ ѓ ӓ ԓ O`2ԓ ѓ jEU1ѓ ԓ CFU1XG* Times ew Romanԓ ѓ PM1ғ ғ =RK1,ғ Cdv% % %  % % % TO2@E@OLahlHistory (Leino H'2"0#<,22% % % Tl2i@E@LahXet al,2TTi2@E@iLahP.% % % T2y@E@Lahp, 2009). The DNA 2232!=2,HIHTz2K @E@zLahpof these 90 to 1402!2,(,232222TTL 2m @E@L LahP-"Tdn 2 @E@n LahTyear0-,"TT 29 @E@ LahP-!T : 2+@E@:  L4&^WMFC<<ahold seeds is severely degraded, 22',,2''',2,!,03-1!,3,2 TXO@E@OWLahbut in spite of this degradation PCR amplification is possible, in particular of fragme22 2 '2, 2! 2' 2,1"-2,22 8CC ,N2!,,12 ' 22''2, 2 2,!,2,! 2! "!-1N,T+@E@ Lah`nts less 2' ,'( TO@E@OrLahxthan 300 bp long (Fig.2,2%222%23%222%"71TT!@E@rLahP &TT"S@E@"rLahP32TlTH@E@Tr0Lah). We have successfully amplified not only multi!%_,%2,3,%'3,,,''!20&-N2!,2%22%220%N3TTIi@E@IrLahP-!Tj+@E@jrLahlcopy genes such ,230&2,2,(%'2,3 TO@E@OLahlribosomal DNA, !22'2N,HHHTt] @E@1Lahbut also single copy genes like microsatellites. 22,'3'21,,2402,2,'2,O,!2',,,'TT^  @E@^ LahP -% % 6h6ah6a66g6`g6`66f6_f6_66e6^e6^66d6]d6]66c6\c6\66b6[b6[66a6Za6Z66`6Y`6Y6 6 _6X_6X 6  6 ^6W^6W 6  6 ]6V]6V 6  6 \6U\6U 6  6 [6T[6T 6 6Z6SZ6S66Y6RY6R66X6QX6Q66W6PW6P66V6OV6O6  .@Times New Roman---@Times New Roman------ F2 _'Barley Genetics Newsletter (2009) 39:20            2 _%-2 _-23 --- 2 _F ,' 2 { ,'2 TW20 2 To ,' 2 q ,'@Times New Roman--- T2  Barley Gen   72 etics Newsletter (2009) 39:20        2 -2 23 2   @Times New Roman--- 2 c P2 ).Historic barley seed samples and their utility           2  --- 2  @Times New Roman------------2 ' M. W. Leino  ---2 81* --- 2 'I #2 'Oand J. Hagenblad    ---2 2,3--- 2  --- 2 8c ---2 T1 ---D2 `+&Swedish Museum of Cultural History, SE          2 `-+2 `643 98 Julita, Sweden     2 ` ---2 q2 ---d2 |;Uppsala University, Department of Ecology and Evolution, SE                2 |!-,2 |)752 36 Uppsala, Sweden     2 | ---2 3 ---2  Norwegian   2  2 Univ.3   2 O 2 Uof  2 i 2 pScie  2 nce 2  2 and 2  2  Technology,   2 h 2 nDep 2  2 of  2  2 Biology,   2   2 N 2 &-,2 .7491 Trondheim, Norway    2 / ,--'a2 19*Author for correspondence: Matti.Leino@nordiskamuseet.seS              2   2   2   2  2 &Gene 2 & 52 &banks around the world hold      2 & over 370.000   2 & ,2 &accessions of barley (    "2 &van Hintum and    2 B Menting, 2003d   D2 B&&). Although a large number of landrace         F2 B's and obsolete cultivars are stored in         2 _]genebanks the geographic coverage of this conserved material is far from evenly distributed.                         2 {XFor example only a handful of landraces from Northern Europe have been conserved (Jones                       2 {#et  2 al,  2  2 2008 2 a /2 ). Thus studies of geogr     +2 aphic differences in o     2  land race  42 barley are hampered by the      >2 "availability of suitable material.           2   2  ;2  Another major drawback with gene        2  X2 3bank material is the authenticity of the available               2 Naccessions. Evolutionary forces such as drift and selection may have affected                2  the genetic   72 %diversity since the accessione       2 %s 2 % 2 %w2 %er  2 %e 2 % %2 %deposited in gene   2 % 2 %bank 2 %s 2 %  2 % and there is     2 %also g  2 % a risk of  )2 Acontamination from p      2 Amoor pr  (2 Aopagation practicesp    2 An 2 Ax (Parzies    2 Aet al.,,   2 A  2 A2000).  2 AM. 2 AS ,2 A\Furthermore, the true       d2 ];geographic origin of cultivars may be unknown or uncertain.                  2 ]  2 z  2 z 2 Re /2 cently material from non        2 -:2 viable historic collections hase           2   2 become ,   2 |recognized as    2 an  82 alternative to the use of gene         2  g2 =bank material. Herbarium material can be well documented and                 2 Stherefore associated with useful geographic information, but the amount of material                       2  2  available   2  is often    q2 Dlimited and successful genotyping can prove problematic (Savolainen                  2 et al.,,   2  2 1995 2  ). In  X2 3contrast, seed collections can contain equally well              2 -G2 (documented samples with large number of            b2 $:seeds from each accession, thus enabling studies on a popu         52 $lation level. Moreover, DNA        2 @_quality of aged seeds widely surpasses that of other equally old vegetative tissue like straw,                      #2 \leaves and chaff    2 \2 2 \8(Lister   2 \et al.,   2 \ 2 \2008).  2 \. 2 \  2 y 2 \In Sweden there are two large seed collections, both assembled at the turn of the last centu                          2 ry. n2 BThe collection at the Swedish Museum of Cultural History contains                     2 C419 2 h 2 p different    2  accessions   @Times New Roman-@Times New Roman- @Times New Roman-@Times New Roman- @Times New Roman--)2 of barley from 1865      2 ^ 2 k 2 q1918 . 2 (Leino   2 et al.,   2 ' 2 . 2009). These   2 samples   "2  were collected      2  on farm    2 from e q2 Dmany different counties across Sweden and Finland with a minor addit                 2 Tion of   2  accessions  2  2 from f  @Times New Roman-@Times New Roman--2  the rest of E   #2 !urope (Fig. 1).    F2 'Another seed collection is stored at V            #2 znersborg Museum    2 " (Johansson 2 ", 2003)   2 "F 2 "M and contains     2 "61  %2 "barley accessions6    L2 "+. The samples in this collection originate               I2 ?)from an agricultural exhibition in 1882. e          2 ?3The seeds are   C2 ?%in both cases stored in sealed glass V       2 [Tcontainers and were displayed on exhibitions and museums but have never regenerated                        2 [(2 [Fig. 3   2 w2 2 w). 2 w  2  2 VWe have genetically analysed part of the collection at the Swedish Museum of Cultural                    @Times New Roman------"2 History (Leino      ---2 *et al3   2 U.---%2 [, 2009). The DNA   &2 of these 90 to 140     2 -2 year  2 -;2  old seeds is severely degraded,         2 Wbut in spite of this degradation PCR amplification is possible, in particular of fragme                          2  nts less    ,2 than 300 bp long (Fig.    2 z 2 3 S2 0). We have successfully amplified not only multi          2 -#2 copy genes such "2 !ribosomal DNA,   U2 !<1but also single copy genes like microsatellites. l              2 !  "System--՜.+,D՜.+,h$ hp  Nordiska museetG' /Historic barley seed samples and their utility Title 8@ _PID_HLINKSA<Pehttp://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Powell%20W%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus  7http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Brown%20TA%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus 2http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Jones%20MK%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus =http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Jones%20G%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus l\http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Charles%20MP%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus ' http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Smith%20LM%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus 8 http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Bower%20MA%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus Tnhttp://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Mackay%20I%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus ;http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Leigh%20FJ%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus =http://www.ncbi.nlm.nih.gov.ezproxy.its.uu.se/sites/entrez?Db=pubmed&Cmd=Search&Term=%22Jones%20H%22%5BAuthor%5D&itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DiscoveryPanel.Pubmed_RVAbstractPlus   !"#$%&')*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRTUVWXYZ[\]^_`abcdefghijlmnopqrstuvwxyz{|}~ Root Entry F )pData (U1TableS9/WordDocument ANSummaryInformation(kDocumentSummaryInformation8pMsoDataStore/@4SY0VX3DB5PUA==2/@Item  PropertiesUCompObj y   F'Microsoft Office Word 97-2003 Document MSWordDocWord.Document.89q