Genetic analysis of rice allelopathy

REPORTS7. Meuwissen, T. H. E., Goddard, M. E, The use of marker haplo-Chinese Science Bulletin 2003 Vol. 48 No.3 265 - 268types in animal breeding schemes, Genetique, Selection, Evolu-Genetic analysis of ricetion, 1996, 28: 161.8. Larzul, C., Manfred, E, EIsen, J. M., Potential gain from includ-allelopathying major gene information in breeding value estimation, Genetics,Selection, Evolution, 1997, 29: 161.ZENG Dali', QIAN Qian', TENG Sheng',). Dekkers, J. C. M., van Arendonk, J. A. M., Optimizing selectionDONG Guojun', H. Fujimoto2, Kunihifo Yasufumi2for quantitative traits with information on an identified locus in& ZHU Lihuang3ourtbred populations, Genetical Research, 1998, 71: 257.1. Key Lab for Rice Biology, Ministry of Agriculture, Institute of China10. Liu Huiying, Zhang Qin, Zhang Yuan, Relative eficiency ofNational Rice Research, Hangzhou 310006. China;marker assisted selection when marker and QTL are incompletely2. Japan Intermational Research Center for Agriculture Science, Tsukuba305-8686, Japan;linked, Chinese Science Bulletin, 2001, 46: 2058.3. Institute of Genetics, Chinese Academy of Sciences, Beijing 100101,11. Georges, M.. Case history in animal improvement: Mapping com-Chinaplx tats in ruminant, Molecular Disetiono of Complx Traits Crrespondence shoud be adrsed to0 Zhu Lihuang (e-mail: lhzhu@genetics.ac.cn) or Qian Qian (e-mail: qianqian@ mail.hz.zj.cn)(ed. Paterson, M. E.), Boca Raton: CRC Press, 1998, 512 - 536.| 2. Ruane, J., Colleau, J. J., Marker-assisted selection for a sex-limit-Abstract A double haploid population derived fromed character in a nucleus breeding population, Journal of dairyanther culture of ZYQ8/JX17, a typical indica and japonicaScience, 1996, 79: 1666.hybrid, was used in this study. The inhibited effect of13. Gomez Raya, L., Klemetsdal, G., Two-stage selection strategieswater-soluble extract of 123 DH pure lines leaves on thelettuce roots growth was investigated, and the QTLs analysisutilizing marker-quantitative trait locus information and individualof rice allelopathy was carried out. Totally, four QTLs relat-performance, Journal of Animal Science, 1999, 77: 2008.ed to rice allelopathy were detected, and they were on chro-14. Schulman, N. F, De Vries, M. J, Dentine, M. R., Linkage dise-mosomes 3, 9, 10 and 12, respectively. The LOD scores werequilibrium in two-stage marker assisted selection, Jourmal of Ani-3.40, 2.68. 2.75 and 3.08, respectively. Among them, additiveeffects of the QTLs on chromosomes 3 and 10 were 1.65 andmal Breeding and Genetics, 1999, 116: 99.15. Falconer, D. s, Mackay, T. F. C., Introduction to Quantitative1.43, on chromosomes 9 and 12 were - 1.44 and -1.58, respec-tively. Allelopathy characteristics of another three commonGenetics, 4th edition, Essex: Longman, 1996.rice varieties were also studied.16. Wang, T, Femando, R. L., van der Beek, s. et al., CovarianceKeywords: rice, alopathy, QTL, biology detection, lettuce.between relatives for a markered quantitative trait locus, Genetics,Selection, Evolution, 1995, 27: 251.Allelopathy refers to the effects of metabolic secre-17. Spelman, R. J., Garrick, D. J, van Arendonk, J. A. M., Uilisationion of plants or microorganism on the environmentl,2of genetic variation by marker assisted selection in commercialPeople found that some crops, such as oats and cucumberdairy cattle population, Livestock Production Science, 1999, 59:had the inhibited effect on the field weedslB.41. Because1.allelopathy utilized plants self-defense and anti-adversity8. Riquet, J., Coppieters, W., Cambisano, N. et al., Fine-mapping ofability and would not lead to the environmental pollutionquantitative tait loci by identity by desent in outbred populations as herbicide did, it had important significance in the dApplication to milk production in dairy cattle, Proc. Natl. Acad.velopment of sustainable and environment- protectiveSci. USA, 1999, 96: 9252.agriculture.19. Thaller, G., Hoeschele, L, Fine -mapping of quantitative trait lociRice allelopathy study started from the germplasmscreening. Chou et al.b5l screened 24 oryza perennis mo-in half- sib families using current recombinations, Genetical Re-ench for allelopathy. Fujii et al.!6] used lettuce as biologi-search, 2000, 76: 87.cal detecting means, allelopathy characteristics study of20. Ruane, J, Colleau, J. J. Marker assisted selection for genetic im-189 Asian and African cultivated rice varieties indicatedprovement of animal populations when a single QTL is marked,that there were significant differences among rice varietiesGenetical Research, 1995, 66: 71.in field weeds controlling. Hassan et a1.(7l found that 3021. Spelman, R. J, van Arendonk, J. A. M., Effect of inccurateand 10 rice materials could control the field Barnyardparameter estimates on genetic response to marker-assisted selec-grass (Echinochloa crusgalli) and Nutgrass flatsedge (Cy-tion in an outbred population, Journal of dairy Science, 1997, 80:perus rotumdus) growth, respectively. Maneechote et al.[8]3399.stud中国煤化工md rice and wild rice on22. Spelman, R.. Bovenhuis, H, Genetic response from marker assist-barnCNMH G: found that a part of themed selection in an outbred population for differing marker bracket hadMYHGrass and lettuce, and thesizes and with two identified quantitative trait loci, Genetics, 1998,inhibited effect on young roots and seedlings was between148: 1389.20% and 60%8]. Kong et al.9 analyzed the differences .(Received August 23, 2002)between the allelopathy and un-allelopathy varieties byChinese S府亦数letin Vol. 48 No. 3 February 2003265REPORTSusing HPLC techniques. On this basis， germplasmtions were done. Samples were cultured at 251( in thescreening and genetic improvement of rice allelopathydark for 3 d and 20 germinated seeds were selected ran-were carried out, and further verified the existence of ricedomly for root length investigation. The allelopathy wasallelopathy9. Reports on genetic mechanism of allelopa-expressed by the lettuce relative root length. The shorterthy were not found at home. In 2001, using the rice water- the relative length was, the stronger the allelopathy wouldsoluble extraction as material and lettuce as biologicalbe. The formula was as follows: Allelopathy = Lettucedetecting means, Ebana et al.(10.11 constructed the riceroot length in extraction liquid/ lettuce root length in bac-allelopathy studying system and developed the allelopathyterium-free water x100%.special rice germplasm screening and genetic analyzing, 7(. i) QTL analysis. Allelopathy characteristics forQTLs on chromosomes 1, 3, 5, 6, 7, 11 and 12, respec-each DH line were transformed by sin-'√p . Based on thetively, which related to rice allelopathy were detected.constructed linkage map of this DH population, intervalRice allelopathy was a new study area to which sci-QTL mapping was conducted by using the software 0entists all over the world paid more attention in recentMapmaker/QTL 1.1 to analyze the QTLs for allelopathydecade. Domestic references on the aspects almost intro-ind PL. The presence of a QTL was determined with aduced aboard studies, and reports on special germpalsmLOD threshold of 2.5. LOD. >2.5 indicated that the high-screening, analyzing, and utilizing of allelopathy were few.In this study, using lettuce as a biological detecting mate-est LOD score position in the interval was just the QTLrial, the inhibited effect of rice water soluble extraction offor the trait. The variation expression and additive effectrice young seedling leaves on the lettuce roots growth wasof each QTL for relative traits were also calculated. QTLinvestigated. Allelopathy of a DH population and its par-nomenclature followed that of McCouch et al.Il2.ents were investigated and genetic analysis was carried2 Resultsout, and the genetic mechanism of rice allelopathy was(.1) Comparison of allelopathy characteristics of DHpreliminary discussed, which provided the basis to screenparents and other rice varieties.Table 1 shows the per-and cultivate special rice germpalsm of inhibiting paddyformance of allelopathy of DH parents and other ricweeds growth.varieties. In the DH parents, the allelopathy of male parent1 Materials and methodsJX17 was stronger than that of the female parent ZYQ8,(_ i) Production of DH population. A typical indicaand their allelopahties were mid-level compared with thevariaty Zhaiyeqing 8 (ZYQ8) and a typical japonicaother three rice varieties. Among the varieties tested,Jingxi 17 (JX17) were used as parents for crossing. The FTaichung Native 1 had the strongest allelopathy and Xi-ushui 11 had the weakest. The effect of extraction of let-anther was isolated and cultured on the induced mediumtuce young root on the lettuce root growth was very weak.SK3. After natural doubled or colchicines treatment, thedouble haploid (DH) plants were obtained.Table 1 Comparison of allelopathy characteristics of DH parents andother rice variceties reported(_ i) Extraction of water-soluble chemical compound.VarietyAllopathyNoteThe method of extraction of water -soluble chemicalcompound was the improvement of Kaworu Ebana' slo. 3Taichung Native 19.0 +0.81 a[10]g young leaves from 10 plants for each DH line wereDular152士0.97 bselected at the six-leaf stage. For common rice varietiesJingxi 1717.4土0.80 b(Dular, Taichung Native 1, Xiu Shui 11 and DH parents),Zhaiyeqing 836.6士0.83 c .the leaf samples were selected from 30 plants after beingXiushui 1178.4 +0.85 d .1)transplanted for 10 d. The treated procedure was as fol-lows: samples were pestled at 0_ (, moved into 50 mLLettuce93.1土3.48centrifugal tube, added with ddH2O 30 mL. The mortara) Data were the average standard. The same letter in the columnwas cleaned 3 times. The cleaning liquid was poured intowas not significant at 0.01 level. b) 3 g lettuce root were extracted at thethe centrifugal tube, mixed evenly, kept at refrigerator forsame condition after 10-d water-culturing, three replications.2 h, vibrated for 1 h, then centrifuged for 15 min (1500r/min). The supernatant liquid was selected and cold(_ i) Performance of allelopathy in the parents andstored until use.DH population. Allelopathy of each DH line showed the(ii) Lettuce culture experiment. 50 lettuce seedscontinuous traits (Fig.1 and Table 2). Allelopathies ofwere placed evenly on the petri-dish covered with filtermos中国煤化工:ents or close to them, butpaper, and 3 mL semi-extraction liquid was added. Thelines:fYHCN M H Gathy also segregated. Forbacterium-free water was used as control. Two replica-1) Yu Linqing, Annual report of China National Rice Research Institute, 2000.266Chinese Science Bulletin Vol. 48 No. 3 February 2003REPORTSexample, DH47, the allelopathy was 1%, almost inhibitedTable 2 Variance analysis of alelopathy in DH linesthe lettuce root growth completely. DH37, the weakest, its .Variance resource DISSMSallelopathy was 59%. Allelopathy showed quantitativeInter- lines122 928 140.08 7607.71 7.691.32 .traits in the DH populations.Intra- linesTotal613which 234 markers were evenly distributed over all 1230rice chromosomes, had been constructed by using thispopulation and it was suitable for QTL analysis. IntervalQTL mapping for allelopathies of water-soluble extractionof young seedling leaves from 123 DH lines was conduct-0ted by using the software of Mapmaker/QTL 1.0. The fourdetected QTLs related to allelopathy distributed on chro-mosomes 3, 9, 10 and 12, respectively (Table 3 and Fig.15452). The additive effects of QTLs on chromosomes 3 andRoot length (% of the contro)10 were 1.65 and 1.43, indicating that the two QTLs fromFig.1. Frequency distrbution of the phenotypic value in DH popula-the allele of JX17 increased the root length, while thetions. Phenotypic value was expressed as the percentage of lettuce rootadditive effects of QTLs on chromosomes 9 and 12 werelength treated with the water-soluble to the control.-1.44 and -1.58, indicating that the two QTLs from the(ii) QTL location. A molecular linkage map, inallele of JX17 reduced the root length.Chr.3Chr. 9Chr: 10Chr.12Dist. /cM MarkerDist., /cM MarkerqAP- 1203.08400乐RG104B39A-CT387RG181CTI2114SO6.90-11 20-2个C1234I5 60--G1106G1751010十B39B9.70 i420-FRG409H -C391B牛CTI05A4.40 FG2SC74A qAP-313 80-19.10-1420-RG4S0 3.40RZ617B3小81.0 RG4635.0小 RG2663607-CD0501S60G10821760 G14813.10-8-iC2521 50NC71111 30--GA223NCT339-G103-G291qAP-1013.70-21 60-202011.10-- G2ISS0.90 大G14411 40--G93FqAP-916 60-C161380S.90RG4825.90 计C3S6A 2.68GAS38.60CT4107.80- HCT2211890-C2235.20. RGS74AT42A24.70-238NC6221380--C746G1931640--GASOS1240-14 30-G249-G29516 8028.20-十-RG4519.20-十RG756G16428- cr21A中国煤化工1.80元GL NG1318YHCNMHGFig. 2. RFLP linkage map showing locations of QTL for allelopathy in rice. Markers are to the right of chromosomes. Solid bar tothe right of chromosomes reprent intervals with LOD - 25, and arrows indicate the position of the peak LOD. The peak LOD score isshown above the solid bar.Chinese S府亦数letin Vol. 48 No. 3 February 2003267REPORTSTable3_ QTLs and parameters related to rice alolathy)yAcknowledgements LU Shufang, Nanjing Agriculture University,Locus ChromosomeMarkerLOD V ariation Additiveparticipated in a part of the study. This work was supported by the Na-intervalscore explained effectqAP-33RG450-- RG2663.40 10.241.65tional Natural Science Foundation of China (Grant No. 33970409) andthe Natural Science Foundation of Zhejiang Province (Grant No.qAP-99CT410- CT62.688.02 -1.44 .300221).qAP-10G2155- C162.758.271.43ReferencesqAP- 12RG181-G1106 3.089.79--1.58.1. Wang, D. L, The review of rice allelopathy, Acta EcologicaTotalb)32.46Sinica, 1998, 18(3): 326.a) QTL nomenclature followed that of McCouch et al. (1997);2. Yan, F, Yang Z. M., Han, L. M., Review on research methods forb) compound QTIL analysis results.allelopathy and allelochemicals in plants, Acta Ecologica Sinica,3 Discussion2000, 20(4): 692.3. Fay,P. K., Duke, W. B., An asessment of allelopathic potentialWeeds management was an important link in ricein Avena Germplasm, Weed Sci, 1977,25: 224.high yielding. Chandlerl13I reported that the rice loss .caused by weeds was about 17% in America, while it was4. Putnam, A. R., Duke, W. B.. Biological suppevidence for allelopathy in accessions of cucumber, Science,as high as 25%- -75% in Thailand, and it also reached1974,185: 370.more than 10% in China. With the increase of direct-seeded rice, weeds control showed more importance. Re-5. Chou, C. H, Chang, F. J. Oka, H. I, Allelopathic potentials ofcently, resources that could obviously inhibit the mainwild rice, Oryza perennis, Taiwania, 1991, 36(3): 201 .paddy weeds such as barnyard grass and Monochoria6. Fuji, Y, The allelopathic effect of some rice varities, in Proceed-vaginals had been found--9.14.151. This laid the basis forings of the International Seminar“Biological Control and Inte-controling the paddy weeds by utilizing the allelopathy.grated Mangement of Paddy and Aquatic Weeds in Asia", October19 - 25, Tsukuba, Japan, 1992, 160.Allelapothy occurred within a long period and was oftenconcealed in the obvious competitions between intraspe-7. Hassan, S. M., Aidy, L R., Bastawisi, A. O. et al, Weed manage-cies or interspecies, and was intervened and disturbed byment in rice using alopathie rice varities in Egypt Paper in theenvironmental factors and microorganisms, all these com-"Workshop on Allelopathy in Rice" , IRRI, Los Banos, Philippines,plicated the allelopathy' s study and utilization.1996.Referring to the method of Ebanal0), allelopathy8. Maneechote, C., Krasaesinhu, P., Allelopathic effects of some up-characteristics of DH population of ZYQ8/JX17 and itsland and wild rice genotypes in Thailand, Paper in the First Worldparents were analyzed in our study. Totally 4 QTLs relatedCongress on Allelopathy: A Science for the Future, Cadiz, Spain,to allelopathy were detected, and the allele from JX17 on .chromosomes 3 and 10 could increase the relative root9. Kong, C. H., Xu, X. H, Hu, F. et al, Using specifi secondarylength, the variation expressions were 10.24% and 8.27%,metabolites as markers to evaluate allelopachic potentials of ricerespectively, while the allele from JX17 on chromosomesvareties and individual plants, Chinese Science Bulletin, 2002,9 and 12 reduced the relative root length. By using the47(10): 839.isolated population of PI3 12777/Rexmont, Ebana detected0. Ebana, K, Yan, W. G., Robert, H. et al, Variation in the allelopao-one QTL related to allelopathy near the marker RG1709thic effect of rice (Oryza sativa L) with water soluble extracts,on chromosome 12, which was in accordance with ourAgronomy Joumal, 2001, 93(1): 12.result. In the DH population, the strongest allelopathy was11. Ebana, K, Yan, W. G., Robert, H. et al, Anaiysis of QIL associ-1% for DH47, which almost could inhibit the lettuce rootated with the allelopathic effect of rice using water-soluble exgrowth completely, the weakest was 59% for DH37. AI-tracts, Breeding Science, 2001 ,51:47.lelopathies of most lines were between the parents or2. McCouch, S. R., Cho, Y. G, Yano, M. et al, Report on QTL no-close to them. In the meantime, certain super parent g-menclature, Rice Genet Newsl, 1997, 14: 11.netic types also existed, showing the quantitative traits.The allelopathy breeding was just the beginning in3. Chandler, J. M., Estamated losses of crops to weeds , in HandbookChina, but the successful use will bring the fundamentalof Pest Mangement in Agriculture (ed. Pimentel, D.), Vol. 1, Bocachange of paddy weeds management, and have importantRaton, FL: CRC Press, Inc, 1981, 95.significance in the future rice production. Genetic analysis,14. Dilday, R. H, Lin, J. Yan, W. G, Ientifcation of alopathy ingermplasm evaluation, and breeding utilization of rice”中国煤化olletion Austar. J Exper. A智allelopathy are key tasks in the future. Because the alle-lopathy is a quantitative trait influenced by multi-genes15.YHC N M H Glclopathic rice for Echinochloaand the detected means are complicated, simplifying thecrus-galli control, In Proc. 2nd Intern. Weed Control Cong, 1996.detected method and utilizing the MAS techniques should1175.be further studied.(Received October 17, 2002)268Chinese Science Bulletin Vol. 48 No. 3 February 2003