PROVEI\ANCE RESEARCH: INVESTICATION OF GENETIC DIVERSITY ASSOCIATED WITH GEOCRAPHY (Chapter 4 of Report FAO/IIIFRO Meeting on Forest Genetics)
Preprinted from Unasylua, Volume 18 (2-3), Numbers 73-74 FOOD AND AGRICULTURE ORGANIZATION OF THE UNITBD NATIONS
Purchased by the Forest Service, United States DeJrartment of Agriculture, for official use 4. Provenance research: investigation of genetrc diversity associated with geography
Ii. Z. C,s.LLaHaN
Sumrnary
Prouenrt,ru)e i'n lorestry relers to the poptilu'tiott of tree,s prehetts'|,^e sulnnTaru ol i,'nformation on genet'ic und' enui- grouinE1 at a prtrticular place ol ori'gin,. Pro"-enance re- ron,mentq,l z-ariabi'l'itu tci,thi'n the ra,n,ge ol the spec'ies. sectrch defines the genet'ic ancl en'uirottmental contponents Objecti,ues and 'procedures ol researclt, shoulcl be u,ell defi,n'ed. questions clesiglnirtg anrl erecut- of Tthenotypic ua,ric.tt'ion os,socioterl with g1eogrrtph'ic source. Manu rnust be consi'rherecl in Infor'ntntion, on proueltance is imynrtrtnt i,n' ctssttr'ing 'ittg a sctmple ol ntang prouena,ttaes to rtccomplish obiectdues. sources of seed, to gi,ue toell-ad'apted,, proilucl'iae trees Biosystemati,c studi,es utili,zing sererul tl'iscipl,i,nes should, anil i'n cli,recting breeclirry ol i,nterraci'a'l anrJ 'interspeci,lic be most effici,ent to dcf.ne patterns ol aari,rt'ti,ott nnd' natural hybri,rls totr:aril acla,ptclti'on to parti'atkt'r lctcal,i,ti,es. Con' bioti,c units hr,tt'ing s'irn,ilur for'm and lu'ttction. Seed cepts of the spec'ies, ol urtrit'tion u:i'thin spe.cies, ol cotttin- source shtd,ies u;ill rhentonstrute ad'aptrtbi'lity, grou'th, und ui,ty in thi,s uu,riation, anrl ol relati'on ol tari,ation to uietd. Theu mcrry also serue as seecl Eources. SeeiJ lactors ol the enui'ronm,ent haue deuelopetl oter the past source stu,il'ies i'n senti,-ctricl lands are lucecl with u,n'ir1ue century. Studie.s ol prouenance shoulcl start u;ith cr coin' Ttroblems.
Chapter 4
'I'he choicc of seed sourccs is onc of thc main factors concepts of genctic divcrsity rvithin species ernerging affecting the establishrnent and productivity of planta- over the past 140 years can also be profitably revierved. tions of forest trees. In thc prescnt practice of silvi- The species, variation rvithin specics, and thc naturc culturc, provenance research provides a sound basis of this variation, rvhethcr it be continuous or discontin- for thc selection of seed sources. Anyone conccrned with uous, all need to be considercd. Finally, considcra- afforestation and reforestation should develop a program tion must be given as to holr to study the grcat divcr- of provcnance research to assist in the selection of secd sity rvithin species. Through the modern biosystematic sources. Such valuable rvork has bcen done in the past. approach patterns of variation can be defined. Through Extensions of this and new research are vitally needed the familiar seed stiurce study the most product'ive prov- on an intcrnational scale because rvithout it future invest- enances can be found. By combining both approach- rnents in afforestation and reforestation rvill not rcturn es the goals of provenance research will be most their maximum revenue. Provenancc research should efficiently attained. be given highest priority at the outset, of anv program of forest tree improvement. Provenance research means rnany things to many MnaNrNe ox' PnovDNANcll begin defining some men. It is, therefore, useful to by Terms referring to that part of thc genetic diversity scope of the of the terms in common usc, outline the rvithin species rvhich is associated rvith geography The subject, and rncntion its general importance. must be defined at the outset to provide a common vocabulary. "Provcnience" and "provenance" arc trvo rvords having the samc meaning. Both refer to R. Z. Crr-r,anrem is Assistant Director, Pacific South- west tr'orest and Range Experiment Station' United States origin or source. They are fully interchangeable, and n'orest Service, Berkcley, California, U.S.A. Other members for consistency only the rvord provenance is uscd here. of the drafting team were P. Bouvarel (France), J. Lacaze Provenance in forestry refers to the particular place (France) and A. I'I6tro (rlo). The auihol also acknowl- trees arc grou'ing or the place of origin of sceds edges the assistancc of his colleagues, partictrlarly Dr. v'here W. B. Critchfield, and the rcvier'v and suggestions of Dr. or trccs. Provenance may refer eithcr to native or to O. Langlet (Sweden). planted trees growing at that place, but its common use is in reference to nat'ive trees. By definition and of tcsting. If juvenile chara,cters can bc uscd to eval- some usage provcnance could also rcft-.r to an individ- uate perforrna,nce, then the time rnny be shortened. ual parent trec. Holever, provenancc should be le- Provenance reserlrch also has inportancc for the stricted to rcfer to the popul:rtion of trees growing at pl:rrrned breeding of hybrids bet'w,een species. As was a place and not, to an individual trce. Ilence, " select- demonstrated in Chapterr 3, crossing species is not ing the proper provenance " means selecting the proper enough. One must direct breeding to producc a hybrid locality. combination of provenances having the highest adapta- Use of the word provenancc in rcference to a region tion and productivity. Duffield (1954) states, " Perform- is improper. This use stems from provenance expcri- ance of hybrids is grcatly influenced by the racial ments. In such experiments trees from one local source origin of the parents ...The spccies hybridizcr cannot rnight differ from all others. In some cases reference escape the provenance problem." has been made to thc broad region represented by one Another inportant aspcct of provenance research source as a provenancc. llore refined research undoubt- is the information provided for breeding h1'brids rvithin edly u.ould show this region to be made up of many species. Hybrids cornbining diffcrent provenances often provenances. combine desirable characters of diffcrcnt races and may Thus, proven&nce has a biological meaning roughly result in hybrid vigor for many characters (Clausen, equivalent to a local population or deme. A prove- Keck, and Heisey, 1948; Wettstein, 1958). Such hy- nance will be some part of a race, ecotype, cline, sub- brids may be adapted to a wider rangc of envirorr- spccies or variety.r mcntal condit'ions than were the parents. With l Spocrns: Scientific name and author Common namc OnrcrN: Country State or province County Latitudc Longitude Elevation .- feet or meters Dctailed location (section, township, range, meridian; direction and distance to town or other landmark) Slope direction Slope steepness _% Forest, type SAF tvne number Site index Soil pH (basic, neutral, acidic) Soil moisture-site condition (dry, moist, t'et) - - Other tree species in stand Cor,r,ncrror: Date of collection Number trccs in collcction Ase -- Average height Average diameter Cross out word I{OT applicable: Collected from: (standing trees) (felled trees) (rodent caches) Trees are in: (plantations) (natural stands) Trees are in: (open) (thin stands) (dense stands) Shedding of secd or fruit: (not started) (starting) (underrvay) (complete) Mcthod of extraction: (air dried) (kiln dried at (crown fcrrm, branching, vigor, disease, insect damage,--oC.) etc.) R,pltenr Date Signatures Collector Address Dealer Forester Freunn 15 - Su,ggested, lorm lor docurnentirtg seerl collect,ions, (Prepared by the Comrnittee on Forost Tree hnprovement,. Society of American Foresters) variables plus a few samples representative of other meager seed production will result in collection from regions might be in order. For biosystematic studies, atypical trees, selection of undesirable alternate loca- thousands of trees from hundreds of locations might tions, and high costs of collection. General or regional be sampled. n'or seed source studies, the size of the failures in seed production doubtless will occur. These sample might be limited to hundreds of trees from require repeated collections in key regions or key local- tens of locations. ities in subscquent years. In this event, long-lived Deciding which specific locations to sample v'ithin seed can be stored in refrigerators without serious loss the desired coverage is most difficult. Primary sampling in viability. If this is done, crop year effects may have should be done parallel to major environmental gra- to be considered in subsequent nursery tests (Callaham dients within taxa. X'or example, samples could be taken and Hasel, 1961). along specified degrees of latitude, on both sides of every major mountain range, and at specified intervals Documentation. The investigator should specify the of elevation, such as 300 to 500 meters. Ideally, at kind and amount of information requircd to describe least 3 to 5 samples should be taken along each and document the origin of his material (Rohmeder, major gradient in the environment,. Secondary sampl- 1962). Forms should be provided for entering the re- ing should complete the picture by filling in samples quired. information. Detailed descriptions u'ill be needed along mountain chains and by bringing in unique of the sample trees and locations (Figure 15). A map populations. These latter populations might be outlying showing the location of each collection should be request- stands or suspected plus or minus stands. ed from the collector. Kind, ol trees. After deciding rvhich locations to Collections by co-operators. To effect an economic sample, the investigator must decide which trees to saving at the outset of a study by making collections sample. A sample representing all classes of trees in through co-operators may be shortsighted. Provenance proportion to their frequency in the stand would show studies are expensive, and they have both long-term the range of natural variability in the population. and large economic implications. The best and recom- This type of collection is neither practical nor desirable. mended procedure is for the investigator to go to the Probably a sample of the dominant and. codominant stands and make the collections himself. Often this trees in a stand, avoiding immediate neighbors, would is impossible; hence thc next best procedure is for him be most satisfactory. Ilowever, this too may be imprac- to organize the work on the ground by personal visits tical if co-operators have to be relied upon to make the to enlist and instruct co-operators. If this is not pos- collections. Seed should never be collected from iso- sible, someone on the ground may arrange for these lated seed trees because of the high chance of self- collections in the same way. pollination. Certainly, tr'ielding's (1962) suggestion to organize Preserving the identity of individual trees through collection services in remote areas js worthy of imple- the collection stage is commendable. This permits bio- mentation. If collection scrvices cannot be provided systematic study of genetic variation within and be- in remote areas, international co-operation in organiz- tween populations. An investigator will not, however, ing expeditions for seed collection should be arranged. be able to retain the identity of many individual trces Only as a last resort should one make collections by when establishing seed source stud.ies in the field. " mail order." If this is necessary, extra precautions must be taken to insure that collectors are properly Number of trees. Ideally, the number of trees to be motivated and instructed to do the required job. Inves- sampled at any locality should fluctuate with the phe- tigators forced to collect via the mails may feel that notypic variation betrveen trees at that locality. More " beggars cannot be choosers." They may obtain far specifically it should fluctuate with the variation of more than expected if they: the most important or most variable trait. If the trees to be sampled were relatively homogenous, perhaps 5 1. give adequate statements of their objectives and to l0 individuals would suffice. If the sample trees justification for their requests; were heterogeneous, 25 to 50 or more might be appro- 2. give-both general and specific requirements with priate. Since the investigator usually does not have an tolerance for adjustment in collections; estimate of the variance between trees at each location, provide data sheets shov'ing the kind of informa- common practice is to collect from l0 to 25 individuals 3. tion and details needed; at each location. In no case should the sample to repre- sent an area be limited to only one or a few trees. As +. provide funds for collection and shipment should far as possible every tree should be equally represented costs be prohibitive for collectors; in each seed lot. 5. acknowledge any and all assistance received prompt- Ti,mi,ng. Collections should be organized at least two ly and profusel5r. months before seed ripening and should be made only in years of abundant seed production. This gives the frrecuti,ng collectiotts. Execution of the sampling collector an opportunity to search for stands and trees design calls for close attention to all the foregoing to meet specified requirements. Collections in years of points. Arrangenents fbr field collections must be completed 'rvell ahcad of seed ripening. Collectors must parisons should give sorne idea of the heritability of' bc located, instructed, trained, and cquipped as required. charactcrs and plasticity of phenotypcs. Particular attention must be given to quarantine restric- Seedlings reprcscnting trecs frorn different prove- tions on the movement of plant materials. Thc pr.in- nances should be grou,n irr a variety of uniform or, cipal investigator should be availnble to ans\{.er ques- preferably, controllcd environments.B This r.vill permit tions from collectors dur:ing the collection period. He an asscssment of hcrita,bility of characters and the inter'- rvill have to considcr and approve alternative procedures action betrveen genotypcs and cnvironrnents. Sorne or locations as necessirry. Processing of collections characters l'hich might be studicd profitably are: rates rnust proceed rapi References Berznn, H. O. Jack p'ine lrom Lake States seed sources Bouvennl, P. L'influence de I'origine des graines d'6pi- 196l d,ifreritl susce,pt'ib.ility to attack b,y the wh,ite 1962 c6a sur la croissance en p6pinidre; Ia pr6cocit6 p,ine rteeu,il. U.S. For. Serv. Lake States For. et Ia frdquence cles pousses d'aotrt. Ann. de It)xpt. Sta., Tech. Note 595. 2 p. l'Tlcole nat,itvruIe d,es eaur et lordts, 19 (3): 415-439. IJonnx, R,. W. Differential frost resistance w-ithin one C.tr-r,eu,ut, R,. Z. Experirncntal l,axonomy: rnore than seed 1958 Eucalgptus species. Aust. J. Sc'i., 2I: 84-86. 1961 source studies. In Recertt ctduances i,n botony. Univ. of Toronto Press, p. 1695-1699. Bouvr.nrr-, Variabilit6 de l'6pic6a (P,icea encelsa Linl<) P. Celr-.r,u.trr, R,. Z. Geographic variability in grou'th of dans franqais: r6pnrtition caractd- 1954 le Jura et 1962 forest trecs. In T.T. Kozlolr-ski, etl., Tree gtrototh. res des divers types. Reu. lor. lranc., 6 (2): Ncw Yorl<, Iionald Prcss, p. 3lf -325. 85-98. C,rr-r--rurrt, R. Z. & Hsnr,, A. A. P'irtus ponderosa: }:'eight, Bouvennr,, P. Lcs repeuplernents artificicls - cons6qLrences 196l growth of wind-pollinated progenies. S'iluae 1958 d'ordre g6n6tiqtre. JoLtr. For\ts Sca., 8/9: 524-535. Genet.. l0: 33-42. l0 C,r.rr-enerr, R. Z. & Lrnnrconr, A. R. Altitudinal varia- JonNssos, II. ItJinigc Fragestellungen der forstlichen 196l t,ion at 20 vears in ponderosa and Jeffrey pines. 1952 Nachkommcnshaftspriifung. Z. Torstgenet., 2z J. For., 59: 814-820. 2_8. C,rlreoe. Fonnsrnv Bn,r.Ncs. Tree brccding. In Annu,atr JonxssoN, H. Utvecklingcn i 15-ariga forsoksodlingar av f 960 report oil forest research,, 11ea,r encling Marclt, 31 , f955 tall i relation till proveniens och odlingsort 1960. Dept. of Northern Affairs & Natural fDevelopment, of l5-year-o]d cultures of pine R,esources, Forcst Research Div., p. 19-20. in relation to pro'i'enance and rrarietyf. Suenslca Skogsu. Fdren,. T,id,slt., 53: 56-88. Cenr,rsr-n, A. Variation in the native Scots pite of Scot- 1955 land. In Report on Eorest Research, lor tlrc year I{.tr,nr-a, A. Zur Synthese der experimentellen Un.tersuch- end,,ing March 195t1. For. Comm., Ct. Britain, 1937 ungen iiber Kli,marassen der Holzartetz. Metsiit. D. 1)a)-l)tr. Tuthimuslait'ol St.r r : r n u tr.- I.'o tr l;s't"[.'n u-o I lrlnov!]ttlJ N T Llonlrrrto u. ll' ctrli - \Vrrrc+u'1, .I. W. Cenoi,l'pic variat,ion in whit,e ash. J. Por. tl).-rJa i.ttg (:o-opet'at';ue sttuly of geoglrapfuia 1,latL. lor 42: 489-495. ,tout'(e.s ol sotLtlrcrtt. pitte seed, ,\cu' Orlea,ns, La., 1941o, p. Li"ti" I,'or. Scrv. SoLrlh. l'orcst JixPi,. Sta. 35 Wrlrcur, J. \Y. Ilcolvpic clifferent,iat'ion in red ash. "I. 1941b For., 12: 591-597. S