CARYOLOGIA Vol. 63, no. 2: 176-183, 2010

Karyological studies of breviscapus and related species

Kexian1,2 Song, Yuehua Wang1, Tingshuang Yi2 and Zhiyun Yang2*

1School of Life Science, University, Kunming 650091, Yunnan, . 2 Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204, Yunnan, China.

Abstract — A karyological study was carried out on 37 populations of Erigeron breviscapus across its distribu- tion region. All populations of the Erigeron breviscapus studied are diploid (2n = 2x = 18). Various chromosome morphologies were revealed, but didn’t show clear geographical pattern. Karyological studies were also fi rstly carried out on two closely related species: Erigeron multiradiatus and Erigeron multifolius, both species are dip- loid with the chromosome number of 2n = 2x = 18. Combining the available chromosome studies, the Erigeron has relatively consistent chromosome diversifi cation with a basic number of 9, and most species contain diploid individuals, which suggested Erigeron at the initial phase of polyploid diversifi cation.

Key words: , chromosome, Erigeron, Erigeron breviscapus, karyotype.

INTRODUCTION DONG 1998). A series of bioactive compounds of fl avonoids and phenoles have been extract- Erigeron L. belongs to tribe Astereae of fam- ed from this herb (GAO et al. 2007; ZHANG et ily Asteraceae. The whole genus contains 390 al. 2007; TAO et al. 2008). It has been received species with cosmopolitan distribution (MAB- wide attention in the last few decades due to its BERLEY 2008). Species of Erigeron are herbs or remarkable pharmacological effects, particular rare subshrubs, usually perennial, rarely annual in neuroprotective effects (LIU et al. 2005), an- or biennial. Thirty-fi ve Erigeron species (13 spe- ticoagulant property (WANG et al. 1992), anti- cies are endemic) are found in China, most of bacterial and antifungal activity (LIU et al. 2003). which are distributed in Xinjiang and south- At present, E. breviscapus has been widely used western China (LIN et al. 1985). Some species of to make pills and injections in China (SUN and Erigeron contain various kinds of chemical com- ZHAO 2009). More than 1000 tons of dry materi- ponents including fl avone, phytosterol, pyro- als of this species are used by medical companies catechol, amino acid and microelements (ZHANG every year in China. The tremendous demand of and ZHANG 2007), which are used to improve E. breviscapus has caused a serious reduction in microcirculation, reduce blood viscosity et al. native populations due to over-harvest. (ZHU et al. 2008 ). Erigeron multiradiatus (Wall.) Benth. is wide- Erigeron breviscapus (Vant.) Hand. - Mazz. ly distributed in alpine and subalpine meadow of is an Chinese endemic species. This species is Qinghai- plateau, , , Ne- one of the most important Chinese traditional pal and Sikkim, at altitude range of 2600-4300m herb, which has been used in Tibetan medicine, (LIN et al. 1985). This has been used in Miao medicine, Yi medicine to treat paralyzed traditional Tibetan medicine to treat various and rheumatism pain for a long time (SHUAI and diseases such as hypopepsia, diarrhea, hepatitis, adenolymphitis, rheumatism and hemiparalysis (DIMAER 1986). Erigeron multiradiatus contains various fl avonoids that are similar to E. brevisca- pus, thus it is usually used as the substitution of *Corresponding author: phone: +86-871-5223136; fax: E. breviscapus. Erigeron multifolius Hand.- Mazz. +86-871-5150227; e-mail: [email protected] is endemic to northwestern Yunnan Province SHORT RUNNING TITLE: KARYOLOGY OF ERIGERON BREVISCAPUS AND RELATED SPECIES 177

Fig. 1 — Sampling localities of E. breviscapus, E. multiradiatus and E. multifolius. The numbers are same as those in Table 1. 178 KEXIAN, WANG, YI and YANG

Fig. 2 — Mitotic metaphase of E. breviscapus (A-G), E. multiradiatus (H) and E. multifolius (I): (A) Lon- glin, Guangxi; (B) Nayong, Guizhou; (C) Baoshan, Yunnan; (D) Xiaozhongdain, Yunnan; (E) Lijiang, Yunnan; (F) Huili, Sichuan; (G) Daguan, Yunnan; H) Muli, Sichuan; (I) Deqin, Yunnan. Bar = 10 µm. and southeastern Tibet of China, which could ples. Karyological and/or chromosome studies be found in alpine and subalpine meadow and have not been carried out on E. multiradiatus forest margins, at altitude range of 2800-4100m and E. multifolius. In this study, we carried out (LIN et al. 1985). Medical usage of this species karyological studies on E. breviscapus across its was not explored. distribution region to detect geographical chro- Karyological studies have been carried out on mosome morphological variation. Chromosome E. breviscapus (FENG et al. 2002; LI et al. 2007). number and karyology were fi rstly studied on E. However, these studies include relative few sam- multiradiatus and E. multifolius. SHORT RUNNING TITLE: KARYOLOGY OF ERIGERON BREVISCAPUS AND RELATED SPECIES 179 1.90 65.56 1.86 65.09 Arm ratio % T.C. 2B 1.64 62.17 2B 1.64 2A 1.78 64.06 2A 1.78 pe asymmetry (KA); Arm ratio = 8m+8sm+2st 3B 66.14 1.95 10m+6sm+2st 2A 1.78 64.01 18/9/2x 10m+8sm 2A 1.63 62.01 1.63 2A 18/9/2x 10m+8sm 090493 18/9/2x 14m+4sm 2A 1.53 60.47 1.53 2A 14m+4sm 090493 18/9/2x 61.79 1.62 2A 10m+8sm 090830 18/9/2x China, Yunnan, Xiaozhongdian; 3280 China, Yunnan, 090624 18/9/2x 14m+4sm 2A 1.53 60.44 1 2 4 Gejiu; 2062 6 China, Yunnan, China, Guangxi, Longlin; 1550 7 8 China, Guizhou, Qingzhen; 1400 9 Xuanwei; 2193 China, Yunnan, 090246 090070 Malong; 2100 China, Yunnan, 090345 Dayao; 2270 China, Yunnan, 2000 Yaoan; China, Yunnan, 18/9/2x 18/9/2x 090381 18/9/2x 090393 6m+4sm+8st 090475 8m+10sm 18/9/2x 090491 8m+10sm 3A 18/9/2x 2A 18/9/2x 2A 2.29 12m+6sm 18/9/2x 1.61 14m+4sm 69.59 1.72 2A 61.64 8m+10sm 10m+4sm+4st 2A 63.19 1.53 2B 2A 1.39 60.41 1.85 1.74 58.18 64.96 63.45 11 13 1967 Yongping; 15 China, Yunnan, 16 Baoshan; 2030 China, Yunnan, Eryuan- -Liantie; 3020 China, Yunnan, 18 Baoshan- -Xiaozhongdian; 2920 China, Yunnan, Eryuan, Niujie; 2260 China, Yunnan, 090501 20 090597 090583 090538 22 090588 18/9/2x 18/9/2x Napa Sea; 3350 24 China, Yunnan, 18/9/2x 18/9/2x 26 18/9/2x 8m+10sm Judian; 1950 6m+10sm+2st China, Yunnan, 27 28 10m+8sm 6m+10sm+2st 2A 090648 Lijiang; 2568 China, Yunnan, 6m+10sm+2st 29 2B 2A Ninglang; 2580 China, Yunnan, 2A 1.91 3A 31 2500 China, Sichuan, Yanyuan; 1.71 090724 18/9/2x 1.93 65.67 China, Sichuan, Huili; 2130 1.67 1.97 33 China, Sichuan, Huidong; 2368 MJ-0879 63.14 65.87 090750 62.56 66.27 35 18/9/2x China, Sichuan, Zhaojue; 2827 090766 12m+6sm 36 18/9/2x 37 Daguan; 1162 China, Yunnan, 090817 38 090796 18/9/2x 2B 6m+12sm 39 Xundian, 2080 China, Yunnan, 18/9/2x Kunming; 2220 090838 China, Yunnan, 6m+12sm 1.56 China, Sichuan, Panzhihua; 1823 18/9/2x 2A 8m+10sm 18/9/2x 090880 4m+12sm+2st 60.91 3A China, Sichuan, Muli 3240 6m+6sm+6st(1SAT) 8m+10sm(1SAT) Deqin; 3480 1.86 China, Yunnan, 18/9/2x 090889 3A 3A MJ-0862 2A 3A MY-292 1.87 65.07 18/9/2x 2.11 2.25 1.70 65.20 1.76 10m+8sm 18/9/2x 18/9/2x 67.84 69.22 18/9/2x 62.94 090698 090775 63.79 10m+8sm 2A 8m+8sm+2st 8m+10sm 2A 6m+12sm 1.70 18/9/2x 18/9/2x 3A 2A 62.98 1.73 3B 1.91 2m+14sm+2st 1.73 4m+14sm 63.32 65.68 2.08 3A 63.35 2B 67.50 2.23 1.96 69.08 66.23 1— Localities and vouchers of plant samples examined, with the results chromosome analysis: karyotype formula (KF); karyoty ABLE sum of the long arms/sum of the short arms; T.C. % = sum of the long arms/total haploid length × 100%; satellite (SAT) sum of the long arms/sum short arms; T.C. Erigeron multiradiatus Erigeron multifolius T species Locality no. Localites and altitudes (m) 3 5 Voucher 2n/basic no./ploidy level KF China, Guizhou, Ziyun; 1500 19 10 China, Guizhou, Nayong; 1900 12 KA 14 090317 Xiangyun; 2090 China, Yunnan, 090364 2190 Yunlong; China, Yunnan, 17 18/9/2x Dali; 2410 18/9/2x China, Yunnan, 21 Jianchuan; 2270 China, Yunnan, 090513 8m+10sm 23 8m+8sm+2st 25 3B 090554 18/9/2x 1880 Yezhi; China, Yunnan, 090591 2.03 Shigu; 2190 China, Yunnan, 2155 Yongsheng; China, Yunnan, 18/9/2x 67.03 30 18/9/2x 32 090712 8m+8sm+2st 34 090726 090739 10m+8sm China, Sichuan, Xichang; 1796 2B China, Sichuan, Meigu; 2313 2B 18/9/2x 18/9/2x Huize; 2250 China, Yunnan, 1.69 62.79 10m+8sm 6m+12sm 090857 2A 090886 18/9/2x 1.78 18/9/2x 64.05 6m+10sm+2st 2B Erigeron breviscapus Erigeron breviscapus 180 KEXIAN, WANG, YI and YANG

MATERIALS AND METHODS number in Table 1. Voucher specimens are de- posited at KUN (Herbarium of Kunming Insti- There are total 37 populations of E. brevisca- tute of Botany, Chinese Academy of Sciences, pus and each population of E. multiradiatus and Kunming, China). E. multifolius were included in this study (Table Somatic chromosomes were studied using 1). The sampling localities were shown in Fig. root meristems from introduced . Actively 1, numbers on the map is same as the sampling growing root tips were cut off and pretreated

Fig. 3 — Karyotypes of E. breviscapus (A-G), E. multiradiatus (H) and E. multifolius (I) : A) Longlin, Guangxi; B) Nayong, Guizhou; C) Baoshan, Yunnan; D)Xiaozhongdain, Yunnan; E) Lijiang, Yunnan; F) Huili, Sichuan; G) Daguan, Yunnan; H) Muli, Sichuan; I) Deqin, Yunnan. Bar = 10 µm. SHORT RUNNING TITLE: KARYOLOGY OF ERIGERON BREVISCAPUS AND RELATED SPECIES 181 18 18 (Rydb.) Cronquist (Rydb.) 18 (Fr.) Schinz & Keller (Fr.) 18 viscidus (A. Gray) Piper 18 philadelphicus M. Pop.* 18 var. var. var. Boriss. 18 acris debilis politus Greene 18 Rydb. ex Porter & Britton Rydb. 18 Greene 18 S.F. Blake S.F. 27, 36 Pursh 18, 35, 36, 45, 54, 63, 80 Greene 18 L. 18, 36, 54 Gaudin 18 Vierh.* 18, 36 18, Vierh.* (A. Gray) A. Gray 18 (Greenm.) G. L. Nesom 18, 36 Ledeb.* 18 Standl. 18 A. Gray 18, 27, 45 Torr. & A. Gray Torr. 18, 27, 36 A. Gray 36 Hoppe & Hornsch.ex Bluff Fingerh. 18, 36 L. 18 Greene 18, 36 Greene 18, (Vierh.) Simmons (Vierh.) 18 Graham 27, 36 Ker Gawl. 18 A. Gray 18 Botsch. 18 Torr. & A. Gray Torr. 18, 36 Greene 18 Viehr 18 Viehr Schur 18 Payson 18 Nutt. 36 subsp. subsp. subsp. orus agellaris E. philadelphicus E. glabratus E. compositus E. canadensis E. unifl E. versicolor E. vagus E. tenuis E. glaucus E. petiolaris E. formosissimus E. humilis E. acris E. acris E. elongatus E. eximius E. basilobatus E. divergens E. borealis E. rusbyi E. fl E. nanus E. oharae E. lassenianus E. modestus E. inornatus E. khorossanicus E. major E. acris E. angulosus E. asper E. seravschanicus E. platyphyllus E. potosinus E. subtrinervis E. purpuratus E. sonnei 18 18 (Greene) Cronquist 18 G.L. Nesom 36 J.K. Henry 18 s Maire 18 (Muhl. ex Willd.) Wagenitz Wagenitz (Muhl. ex Willd.) 27 Hulten 18 glaber formosissimus callianthemus glabratus Gray 18 strigosa var. var. var. var. muirii DC. 27, 36 Scop. 18 (A. Gray) A. Nelson 18 Michx 18 lamprocaulis Botsch. 18 mesatlanticu Tatew. & Kitam. Tatew. 18 Retz. 54 Nutt. 18, 54 Nutt. 18,27 subsp. subsp. A. Gray 18 debilis Muhl. ex Willd. Muhl. ex Willd. 18, 27, 36 Hook. 36 Vierh.* Vierh.* 18 Nutt. 18 A. Gray 36 (Bunge) Botsch. 41, 54 Nutt. 18 var. var. Greene 18 Porter 18 subsp. L. 18, 27 Br.-Bl. 18 Br.-Bl. G.W. Douglas & Packer G.W. 18 (Greenm.) G.L. Nesom 18 (L.) Torr. & Gray (L.) Torr. 18 Worosch. Worosch. 18 Greene 18 subsp. (Hook.) Greene 36 var. var. L. subsp. 18 accidus E. glabellus E. chiangii E. utahensis E. vernus E. philadelphicus E. glabratus E. griseus E. elatus E. eriocalyx E. coulteri E. caespitosus E. tilingii E. radicatus E. sumatrensis E. ochroleucus E. formosissimus E. peregrinus E. hyssopifolius E. karvinskianus E. mairei E. miyabeanus E. acris E. fl E. acris E. acris E. alpinus E. annuus E. aureus E. bigelovii E. salishii E. pinnatisectus E. pumilus E. muirii E. koraginensis E. simplex E. strigosus E. polymorphus taxa from chromosome number databases (Missouri Botanical Garden) 18 18 Erigeron . B. Boivin 18 (Rydb.) Cronquist (Rydb.) 18 (Widder) Huber (Widder) 18 (Cronquist) G.W. Douglas (Cronquist) G.W. 18 A. Gray 18 (Greene) Cronquist 18 Hiitonen 18 (Greene) Jeps . provancheri pallens DC. 18, 27 bellidiastrum candidus conspicuus glabratus subsp Hayata* 18 Hook.* 18 var. G.L. Nesom 18 decoloratus Greene 18 Hook. 36 species are distributed in China var. hartwegii plantagineus G.L. Nesom 18 Nutt. 18 var. var. subsp. H.M. Hall 18 Botsch. 18, 36 Michx. 18 (Banks ex Pursh) Greene 18 Botsch.* 18 Halocsy 18 var. inornatus var. 18 asteroids Kerner 18 subsp. (Lindl.) DC. 18 Cronquist 18 Dittrich 18 Brugger 18 Blake 54 var. (Hook.) Nutt. 18 Emberger & Maire 18 (L.) Pers.* 18, 26, 27 L. 18 orus var. var. Fr. Fr. 18 Vill. Vill. 18 2 — Chromosome numbers of 111 var. var. subsp. L.* 18, 36 lifolius Erigeron ABLE E. gaudinii E. celerieri E. turnerorum E. philade E. glabratus E. grandifl E. epiroticus E. cabrerae E. corymbosus E. veracruzensis E. thunbergii E. neglectus E. peregrinus E. foliosus E. hyperboreus E. inornatus E. lonchophyllus E. metrius * E. acris E. fi Taxon 2n Taxon 2n Taxon 2n Taxon 2n Taxon T 2n Taxon E. acer E. acris E. aequifolius E. annuus E. atticus E. eatonii E. bellidiastrum E. rydbergii E. philadelphicus E. pulchellus E. purpuratus E. subtrinervis E. morrisonensis E. komarovii E. silenifolius E. speciosus E. politus 182 KEXIAN, WANG, YI and YANG with p-dichlorobenzene at room temperature for type 2B with the T.C. % of 66.23. This is the fi rst 2h, then fi xed in fresh Carnoy’s fl uid (absolute report on chromosome number and karyology ethanol: glacial acetic acid = 3:1) at 4°C for ca. 30 of this species. min. Before staining, the materials were hydro- The chromosome numbers of 111 Erigeron lyzed with 1N HCL for 10s at 60°C. Finally, sam- taxa are available in chromosome number da- ples were stained with a drop of carbolfuchsin tabases (Missouri Botanical Garden, http://mo- and squashed. Permanent slides were made using bot.mobot.org/W3T/Search/ipcn.html). The ta- the neutral balsam as the mounting medium. The xa names and chromosome numbers were listed best metaphase plates were photographed with a in Table 2, the distribution information and digital camera (LEICA DM1000) mounted on a references were not included for saving spaces. LEICA DFC490 microscope. The nomenclature The chromosome number of nine Erigeron spe- used for the description of the chromosome mor- cies distributed in China are reported (sign by * phology followed LEVAN et al. (1964). Karyotype on taxa names). Combining with three species asymmetry was determined by the categories of studied here, chromosome numbers of total 114 STEBBINS (1971). Centromeric terminalization Erigeron taxa are known. value (the abbreviation, T.C., T.C. % = sum of the long arms/total haploid length ×100%) was determined according to ARANO (1963). DISCUSSION

Karyotype diversity of E. breviscapus - All 37 RESULTS studied E. breviscapus populations are diploid, which is largely consistent with the previous All three Erigeron species are diploid studies by FENG et al. (2002) and LI et al. (2007). (2n = 2x = 18). Metaphase chromosomes of sev- However, we failed to found triploid individu- en representative populations of E. breviscapus al, which has been reported from populations and each of E. multiradiatus, E. multifolius were of Cangshan and Yangjiaping in Yunnan prov- shown in Fig. 2. Detailed descriptions of the cy- ince (LI et al. 2007). Although wild tetraploid tological features of each species are as follows: E. breviscapus individuals were not found, ar- 1. E. breviscapus - All populations are dip- tifi cial tetraploid individuals were produced in loid with 12 different karyotype formulae (Ta- lab (MEI et al. 2009). Chromosome morphology ble 1). Among them, 2n = 2x = 8m + 10sm (Fig. varied among different populations and twelve 2: A; Fig. 3: A) and 2n = 2x = 10m + 8sm (Fig. 2: different karyotype formulae were detected. G; Fig. 3: G) are the main karyotype formulae. Among them, the main karyotype formulae are Others are 2n = 2x = 8m + 8sm + 2st (Fig. 2: B; 2n = 2x = 8m + 10sm and 2n = 2x = 10m + 8sm Fig. 3: B), 6m + 10sm + 2st (Fig. 2: C; Fig. 3: C), with the percentage of 21.62% and 18.92%, 14m + 4sm (Fig. 2: D; Fig. 3: D), 6m + 12sm (Fig. respectively. However, the variation of karyo- 2: E; Fig. 3: E), 6m + 6sm + 6st (Fig. 2: F; Fig. 3: type formula, karyotype asymmetry, and T.C. % F), 6m + 4sm + 8st, 12m + 6sm, 10m + 4sm + 4st, values did not show clear geographical pattern. 10m + 6sm + 2st, and 4m + 12sm + 2st. Their arm Most populations of E. breviscapus have a karyo- ratios (sum of the long arms/sum of the short type asymmetry of type 2A with arm ratio is less arms) are from 1.39 to 2.29. The asymmetry of than 2.00, indicating that these populations have the karyotype is categorized as 2A, 2B, 3A and relatively low interchromosomal asymmetry and 3B, accounting for 54.1%, 18.9%, 18.9% and a monomodal karyotype. But the type 3A and 3B 8.1%, respectively. The values of T.C. % are is 27% and mainly distributed in Sichuan prov- all over more than 60% except one population ince, which reveals E. breviscapus also evolve to (Malong, Yunnan) with 58.18%. asymmetry, especially in Sichuan. There are a few 2. E. multiradiatus - The karyotype formula points need to mention: the southernmost popu- of 2n = 2x = 18 = 2m + 14sm + 2st (Fig. 2: H; Fig. lation Gejiu (Yunnan) has the special karyotype 3: H). Its arm ratio is 2.23. The karyotype asym- formula, 2n = 2x = 6m + 4sm + 8st, and with the metry is of type 3A with the T.C. % of 69.08. highest arm ratio and T.C. %. Population Yany- This is the fi rst report on chromosome number uan (Sichuan) also has an unique karyotype for- and karyology of this species. mula, 2n = 2x = 4m + 12sm + 2st, the asymmetry 3. E. multifolius – The karyotype formula of of type 3A, and the arm ratio also exceeds 2.00. 2n = 2x = 18 = 4m + 14sm (Fig. 2: I; Fig. 3: I). Its Satellites are found in only two populations arm ratio is 1.96. The karyotype asymmetry is of (Huili and Huidong) in Sichuan Province. SHORT RUNNING TITLE: KARYOLOGY OF ERIGERON BREVISCAPUS AND RELATED SPECIES 183

Diversity of chromosome numbers of Genus LI L., DANG C.L. and HUANG R.F., 2007 — Discovery Erigeron - The big genus Erigeron contains ca. of Triploid Erigeron breviscapus (Compositae) and 390 species widely distributed in the world. its potential for breeding. Acta Botanica Yunnanica, However, this genus has one basic chromosome 29: 38-42. LIN R., CHEN Y.L. and SHI Z., 1985 — Flora Reipublicae number (x = 9) based on available chromosome Popularis Sinicae, vol. 74. Science Press, Beijing. information, which is consistent with 9 is one of LIN Y.R., 1997 — On the primary study of the systemat- the major basic number for family Asteraceae ics and fl oristics of compositae from China. Bulletin (LIN 1997). Diploid is the main ploidy level in of Botanical Research, 17: 6-27. Erigeron, diploid was found in 100 taxa, with LIU H., YANG X.L., DING J.Y., FENG Y.D. and XU H.B., which diploid is the sole ploidy level in 82 taxa, 2003 — Antibacterial and antifungal activity of and both diploid and polyploid were found Erigeron breviscapus. Fitoterapia, 74: 387-389. from other 18 taxa. Triploid are quite common, LIU H., YANG X.L., REN T., FENG Y.D. and XU H.B., 2005 — Effects of Erigeron breviscapus ethanol ex- which is detected in 11 taxa. Triploid is usually tract on neuronal oxidative injury induced by super- produced by the unreduced diploid gamete and oxide radical. Fitoterapia, 76: 666-670. haploid gamete, and Triploid plays an impor- MABBERLEY D.J., 2008 — Mabberley’s Plant-book. A tant role in formation of polyploid (RAMSEY et Portable Dictionary of Plants, Their Classifi cations al. 1998). Multiple triploids in gene Erigeron and Uses. Cambridge University Press, New York. suggested this genus at beginning of polyploidi- MEI L., ZHENG S.X., YANG S.C., GUAN W.L., ZHENG zation evolution. Polyploidy is relatively rare X.Y. and LI Y.F., 2009 — Study on morphology and cytology of Erigeron breviscapus plants with differ- in Erigeron, and only seven taxa has ploidy ent ploidy. Guizhou Agricultural Sciences, 37: 134- level equal or higher than tetraploid. Together 135. with three chromosome report in this study, 12 RAMSEY J. and SCHEMSKE D. W., 1998 — Pathways, Erigeron taxa distributed in China with available mechanisms and rates of polyploidy formation in chromosome numbers, nine species are diploids fl owering plants. Annual Review of Ecology and and other three species have both diploid and Systematics, 29: 467-501. polyploid chromosome numbers. SHUAI J. and DONG W.W., 1998 — Experimental re- search of PKC inhibitor, Erigeron breviscapus on the Acknowledgements — The authors are indebted ischemic/reperfusional brain injury. Chinese Phar- macology Bulletin, 14: 75-77. to Professor XUN GONG for his valuable help and to STEBBINS G.L., 1971 — Chromosomal evolution in Mr. SHU-DONG ZHANG for his help during the col- lection of samples (Kunming Insititute of Botany). higher plants. Edward Arnold Ltd., London. This study was supported by the Fund of State Key SUN H.D. and ZHAO Q.S., 2009 — A drug for treat- Laboratory of Phytochemistry and Plant Resources ing cardio-cerebrovascular diseases — phenolic com- in West China, Germplasm Bank of Wild Species pounds of Erigeron breviscapus. Progress in Chem- and CAS Innovation Program of Kunming Institute istry, 21:77-83. (grant no. 540806321211). 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