FEATURE Hawthorn () Resources in China Taijun Guo and Peijuan Jiao Institute of Special Wild Economic Animals and , Chinese Academy of Agricultural Science, Zhuojia, Jilin Province 132109, People’s Republic of China

Fruit of the genus Crataegus are used for eastern and northwestern regions where eco- sessed distinctive botanic characteristics and fresh consumption and processing and as in- logical environments are harsh. For example, isoperoxidase bands in comparison with the gredients in Chinese medicines. In China, they C. pinnatifida is widely distributed in regions common C. pinnatifida (Feng and Guo, 1986; are regarded as “fruits for good health.” China between the Qinling Mountains and the Guo et al., 1989, 1991, 1992). It can be used as is one of the major sites of origin of Crataegus Heilongjiang River and its distribution covers a parent for cold hardiness and polyploid breed- species and has a long history of hawthorn wide geographic and ecological environments. ing. cultivation. According to “Erya,” the histori- Crataegus pinnatifida has considerable varia- Crataegus scabrifolia is the hawthorn cal records written by an unknown author in tion in chromosome numbers and is worthy of mainly in the Yunnan-Guizhou Plateau. Fea- 600 B.C., hawthorn was called “Qiu.” At about further study. tures are tree ≈8 to 15 m high; fruit (diameter A.D. 1000, cultivation started because the 1.5 to 3.2 cm) oblate, yellowish-brown, green need for hawthorn as medicine and preserva- Variation in biochemical composition with red patches; flowering period late March tion increased. Systematic studies on haw- to early April; ripening time mid-August to thorn resources in China started in the 1950s. Fruit of all 12 species (Table 2) contain late September; corymb or compound corymb Since the 1970s, the Institute of Special Wild quercetin, hyperosid, citric acid, methylesters, with 8 to 20 flowers in each inflorescence, Economic Animals and Plants, Chinese Acad- (-)-epicatechin, flavan polymers, and sucrose. three to five styles, 20 stamens; SSC 4.0% to emy of Agricultural Science, in cooperation Crataegus pinnatifida and C. cuneata had 10.8%; acid 1.7% to 3.0%; vitamin C 18 to 97 with other institutes, has surveyed and studied chlorogenic acid but not rutin in their fruit mg/100 g fresh fruit; total flavonoids 18 to 68 hawthorn resources in 22 provinces and mu- (Xie, 1984; Yen, 1982). Crataegus dachurica mg/100 g fresh fruit. nicipal regions in China. This paper summa- and C. maximowiczii did not contain either , an excellent dwarfing rizes hawthorn resources in China based on chlorogenic acid or rutin. The fruit of the rest rootstock (Wan and Cai, 1986), is a shrub ≈1.0 these surveys and studies, as well as our re- of the species listed (Table 2), except C. to 2.0 m high; fruit (diameter 1.0 to 2.0 cm) search in recent years. shensiensis, contained rutin but not chloro- spheroidal or oblate, red or yellow; flowering genic acid. Therefore, rutin and chlorogenic period May to June; ripening time September I. WILD HAWTHORNS acid possibly could be used as indicators for to October; corymb with five to seven flowers classification of Crataegus. Dong (1989) iso- in each inflorescence, four to five styles, 20 Species and distribution lated apigenin-8c-glucosylucoside from haw- stamens; SSC ≈7.5%; acid ≈2.1%; vitamin C thorn leaves and found that flavonoid content ≈31 mg/100 g fresh fruit. Crataegus pinnatifida The genus Crataegus has more than 1000 in the leaves changed with the seasons. Types scions grafted on C. cuneata rootstock were species worldwide. Sixteen species and five and concentrations of flavonoids in the leaves able to fruit in 2 years and were only 1.8 m high botanical cultivars have been identified and differed from those in the fruit. Leaves con- when they were 8 years old, with an average confirmed in China (Nie and Ren, 1982; Wang tained primarily vitexin, rutin, and quercetin, yield of 55 kg/tree. Crataegus cuneata also and Yang, 1993; Xin and Zhang, 1984; Yu, while fruit mainly contained (-)-epicatechin had good graft compatibility with other haw- 1977). The species listed in Table 1 are distrib- and rutin. Flavonoid contents in leaves were thorns when it was used as a rootstock. Trees uted among 26 provinces and autonomous significantly higher than those in fruit (Liu, on C. pinnatifida rootstock were twice as tall regions in China. Among the species, C. 1990). Hawthorn fruit contain vitamins B1, as those on C. cuneata. pinnatifida Bge. is most widely distributed, B2, B6, and C, and 17 amino acids (Liu, 1990). Crataegus hupehensis, containing impor- from the Chinling Mountains to the The total content of amino acids in the fruit tant ingredients in Chinese medicines, is a Heilongjiang River (Beijing China Institute of was 3.1%. tree ≈3 to 4 m high; fruit (diameter 1.0 to 2.5 Botany, Academia Sinica, 1974). This species cm) spheroidal, red or yellow; flowering pe- has three botanical cultivars. Crataegus Species with unique characteristics riod May to June; ripening time August to pinnatifida usually has red fruit; however, September; corymb with 20 to 25 flowers per yellow-fruit types also were found in Shanxi Six of the hawthorn species in China (Table inflorescence, five styles, 20 stamens; SSC and Shandong provinces. In our breeding pro- 1), C. pinnatifida Bge., C. scabrifolia (Franch.) 3.8%; acid 2.1%; vitamin C ≈29 mg/100 g gram, we noticed that even if both parents of C. Rehd., C. cuneata Siebet Zucc., C. hupehensis fresh fruit. pinnatifida cultivars were red-fruit types, their Sargent, C. altica (Loud.) Lange, and C. Crataegus altica and C. kansuensis seeds

F1 generation would have some yellow-fruited kansuensis Wils., are more important than the were capable of germinating normally after 3 individuals. This characteristic is very impor- other 10 (Zhang et al., 1982). to 4 months of stratification, 1 year earlier tant in studying C. pinnatifida’s origin, evolu- Crataegus pinnatifida is the major species than the seeds from other Crataegus species. tion, and genetics. of hawthorn and rootstock north of the Yellow In China, C. altica is a tree ≈6.0 m high; fruit River. Characteristics are tree ≈6.0 m high; (diameter 0.8 to 1.0 cm) spherical, golden; Chromosome numbers fruit (diameter 1.0 to 3.6 cm) pyriform or flowering period mid-May to June; ripening globular, red or yellow; flowering period late time mid- to late August; compound corymb The chromosome numbers of eight species May to early June; ripening time, early August with 12 to 68 flowers per inflorescence, four we observed and four species reported by to early October; corymb with 14 to 25 flowers to five styles, 20 stamens; SSC ≈9.2%; acid others were either 34, 51, or 68 (Table 2). per inflorescence, three to five styles, 20 sta- ≈0.6%; vitamin C ≈29 mg/100 g fresh fruit; Among the 12 species listed, five are poly- mens; soluble sugar concentration (SSC) pectin ≈1.9%. Crataegus kansuensis is a shrub ploids or have more than one type of ploidy. ≈8.5%; acid ≈3.5%; vitamin C 20 to 118 mg/ or tree ≈2.5 to 8.6 m high; fruit (diameter 0.8 The polyploid species or species with poly- 100 g fresh fruit; total flavonoids 0.4% to to 1.0 cm) spherical, red or orange; flowering ploidy were distributed mainly in the north- 0.8%. period mid-May; ripening time mid- to late A unique polyploid group of C. pinnatifida August; corymbs with eight to 18 flowers was found on the Changbai Mountain, Jilin per inflorescence, two to three styles, 15 to Received for publication 9 Nov. 1994. Accepted for Province. It was characterized by pollen abor- 20 stamens; vitamin C 23 to 32 mg/100 g publication 24 Mar. 1995. tion, early ripening, and large fruit, and pos- fresh fruit.

1132 HORTSCIENCE, VOL. 30(6), OCTOBER 1995 Table 1. Distribution of Crataegus species and their botanical cultivars in China. Species Province or autonomous region C. scabrifolia (Franch) Rhd. Yunnan, Guizhou, Sichuan, Guangxi C. hupehensis Sarg. Hubei, Hunan, Jiangxi, Jiangsu, Zhejiang, Sichuan, Shaanxi, Shanxi, Henan C. cuneata Siebet Zucc. Henan, Hubei, Jiangxi, Anhui, Hubei, Jiangsu, Zhejiang, Yunnan, Guizhou, Guangdong, Guangxi C. cuneata var. longipedicellata M.C. Wang Fujian, Shanxi C. wilsonli Sarg. Henan, Hubei, Shaanxi, Gansu, Zhejiang, Yunnan, Sichuan C. kansuensis Wils. Gansu, Shanxi, Shaanxi, Guizhou, Sichuan C. shensiensis Pajark. Shaanxi C. pinnatifida Bge. Heilongjiang, Jilin, Liaoning, Inner Mongolia, Hebei, Henan, Shandong, Shanxi, Shaanxi, Jiangsu C. pinnatifida var. major Br. Heilongjiang, Jilin, Liaoning, Inner Mongolia, Hebei, Henan, Shandong, Shanxi, Shaanxi, Jiangsu C. pinnatifida var. psilosa Schneid. Heilongjiang, Jilin, Liaoning, Inner Mongolia, Hebei, Henan, Shandong, Shanxi, Shaanxi, Jiangsu C. pinnatifida var. geholensis Schneid. Liaoning C. oresdbia Smith. Yunnan C. maximowiczii Schneid. Heilongjiang, Jilin, Liaoning, Inner Mongolia C. maximowiczii var. ninganensis Nie. Heilongjiang C. sanguinea Pall. Liaoning, Jilin, Heilongjiang, Hebei, Inner Mongolia C. dahurica Koehne. ex Schneid. Heilongjiang, Inner Mongolia C. aurantia Pojark. Shanxi, Shaanxi, Gansu, Hebei C. chungtienensis Smith. Yunnan C. altica (Loud) Lange. Gansu, Shanxi, Shaanxi, Guizhou, Sichuan C. remotilobata H. Raik. Xinjiang C. songarica K. Koch. Xinjiang

Table 2. Chromosome number of 12 Crataegus Variation in chromosome numbers Summary species in China. Among 63 cultivars we observed (Guo et The research program on hawthorn re- Chromosome al., 1989) and 34 cultivars reported by others sources in China is aimed at selecting com- Species no. (x = 17) Referencez (Pu et al., 1987; Song and Lin, 1985; Zhang mercial cultivars suitable for fresh consump- C. scabrifolia 2n = 2x = 34 3 C. hupehensis 2n = 2x = 34 3, 4 and Xin, 1986), 16 were polyploid (2n = 3x = tion, processing, and ingredients of Chinese C. cuneata 2n = 2x = 34 2, 3, 4 51 and 2n = 4x = 68), which was 16.5% of all medicines. The ideal cultivars are the ones C. wilsonii 2n = 2x = 34 1, 2, 3 observed cultivars. These polyploidies were with large fruit, high nutrient contents, and C. kansuensis 2n = 2x = 34 1, 3 mainly distributed in cold Northeast China high concentrations of medical ingredients. C. shensiensis 2n = 2x = 34 2 and in sunny, warm Shandong Province. Poly- Besides the Institute of Special Wild Eco- C. dachurica 2n = 2x = 34 1, 4 ploidies grow vigorously and have higher nomic Animals and Plants, Chinese Academy C. pinnatifida 2n = 2x = 34 3, 4 adaptability. All hawthorn cultivars in China of Agricultural Science, a few other organiza- 2n = 3x = 51 1, 2 have been selected from native stands. tions also are involved in research on haw- 2n = 4x = 68 1, 2 thorn resources in China. Examples are C. maximowiczii 2n = 2x = 34 1, 4 2n = 4x = 68 2 Variation in ripening time Shenyang Agricultural Univ. in Shenyang, C. sanguinea 2n = 3x = 51 4 Liaoning Province, and Beijing Academy of 2n = 4x = 68 1 Hawthorn cultivars varied dramatically in Agricultural Science in Beijing. For more in- C. songarica 2n = 4x = 68 2 their fruit ripening time (Table 3). Fruit of formation on hawthorn research and commer- C. altica 2n = 4x = 68 1, 3 early ripening cultivars needed only 90 days to cial production in China, readers are referred z1 = Guo et al., 1989; 2 = Pu et al., 1987; 3 = Song mature, while 180 days were needed for late- to two reference books, which will be pub- and Lin, 1985; 4 = Zhang and Xin, 1986. ripening cultivars. Generally, early ripening lished soon in China. They are Encyclopedia cultivars had smaller fruit, could not be stored of Fruit Crops in China–Hawthorn and De- for long, and were difficult to transport. How- scriptor List for Hawthorns (Crataegus). ever, fruit of early ripening cultivars contained II. HAWTHORN CULTIVARS appropriate quantities of sugar and acid, mak- Literature Cited ing them suitable for fresh consumption. China has more than 1000 years of history Beijing China Institute of Botany, Academia Sinica. of cultivating hawthorns. However, to our Variation in fruit sizes 1974. Flora. vol. 36 (in Chinese). Science Press knowledge, there was no report on hawthorn China, Beijing. p. 189–206 cultivar resources until the 1950s. Wen (1960) Fruit size varied significantly from cultivar Cao, Z. 1983. Hawthorn cultivar: Liaohong. China reported some local hawthorn cultivar re- to cultivar (Table 3). Average fruit length fruits (in Chinese). 1:22. sources in Shandong and Liaoning Provinces. ranged from 1.5 to 2.9 cm and fruit diameter Dong, Y. 1989. Study of flavonoid composition of Since the 1970s, many good cultivars have from 1.6 to 3.6 cm. The average weight of 100 hawthorn leaves (in Chinese). MS Thesis, Jilin Agr. Univ., Changchun. been selected and are being used in commer- fruit ranged from 340 (‘Ji Fu Yi Hao’) to 1600 Feng, B. and T. Guo. 1986. Study on pollen abortion cial production. g (‘Da Jin Xing’). Cultivars used by farmers and GA-induced parthenogenesis of early-rip- Valuable cultivars or types have been found were mainly the ones with large fruit and were ening hawthorn (in Chinese). J. Spec. & Valu- in C. pinnatifida, C. scabrifolia, C. hupehensis, late ripening. able Wild Animals & Plants 3:11–14. C. cuneata, C. sanguinea, and C. dahurica. Fu, L. 1989. Study of hawthorn resources of Jilin However, hawthorn cultivars in China have Variation in fruit colors Province (in Chinese). MS Thesis, Jilin Agr. basically originated from three species: C. Univ., Changchun. pinnatifida, C. scabrifolia, and C. hupehensis. Colors of hawthorn fruit can be grouped Gu, M. and D. Shen. 1987. Breeding and improve- About 150 cultivars used in the major haw- into two categories: red and yellow. Within the ment of several major fruit crops in China (in Chinese). Jilin Agr. Sci. 3:68–74. thorn production areas are mainly from C. red group, color differences of fruit flesh be- Guo, T., B. Feng, and X. Sun. 1989. Observation on pinnatifida. Fruit botanical (Table 3) and tween cultivars were relatively large. Flesh chromosome numbers of hawthorn cultivars in chemical (Table 4) characteristics of cultivars color can be divided into three categories: red, China (in Chinese). J. Spec. & Valuable Wild of this species, which are primarily distributed pink-white, and green (Cao, 1983; Zhao, 1979). Animals & Plants 3:14–16. north of the Yellow River, vary widely. Farmers mainly use red-fleshed cultivars. Guo, T., P. Jiao, J. Li, B. Feng, and X. Sun. 1991.

HORTSCIENCE, VOL. 30(6), OCTOBER 1995 1133 FEATURE

Table 3. Fruit characteristics of major hawthorn cultivars of Crataegus pinnatifida. Fruit size (cm) 100-Fruit Fruit coat Chromosome Ripening Fruit flesh Cultivar Length Diam wt (g) colorz no. timey colorx Da Wang 2.4 2.2 625 DR 2n = 3x = 51 L/September PiW-PiR Ye He 2.4 2.5 625 DR 2n = 2x = 34 E/October PiW-PiR Ji An Zi Ro 2.3 2.5 833 LPR 2n = 2x = 34 E/October PuR Xi Feng Hong 2.5 2.8 900 PR 2n = 2x = 34 E/October DR-PuR Liao Hong 2.5 2.8 769 DR 2n = 2x = 34 E-M/October DPi-PuR Yan Yang Hong 2.4 2.5 768 DR 2n = 2x = 34 E/October DPi-DR Yan Yang Qing 2.5 2.7 830 DR 2n = 2x = 34 E-M/October PuR-DR Jin Xing 2.7 2.7 980 LR 2n = 2x = 34 E-M/October GW Han Lu Hong 2.5 2.6 769 DR 2n = 2x = 34 E-M/October PiW-PiR Mian Zha 2.5 2.8 830 R 2n = 2x = 34 E-M/October GW-PiW Hong Yang Mian 2.7 3.6 1200 DR 2n = 3x = 51 L/October YW Bai Yang Mian 2.5 3.2 1200 DR 2n = 3x = 51 L/October PiW-PiR Chang Ko 2.7 3.2 1000 DR 2n = 3x = 51 L/October GW Da Jin Xing 2.9 3.5 1600 DR 2n = 3x = 51 L/October GW-PiW Yu Bei Hong 2.6 2.8 1000 R 2n = 2x = 34 M/October PiW-PiR Yan Guo Hong 2.8 2.6 870 LPR 2n = 2x = 34 E/October PiR-DR Qu Jin Xing 2.0 2.1 560 DR 2n = 2x = 34 L/September PiR-DR Shuang Hong 1.9 2.3 500 LR 2n = 2x = 34 L/September PiR-DR Fu Li Hong 1.5 1.6 362 LR 2n = 3x = 51 E-M/August PiW Ji Fu Yi Hao 1.9 2.0 340 PR 2n = 3x = 51 E-M/August PiR Zuo Fu Yi Hao 1.7 2.1 355 LPR 2n = 4x = 68 L/August PiR zD = dark, L = light, P = purple, and R = red. yE = early, L = late, and M = middle. xD = dark, G = green, Pi = pink, Pu = purple, R = red, W = white, and Y= yellow.

Table 4. Fruit biochemical composition of major hawthorn cultivars of Crataegus pinnatifida. cultivars of Chinese hawthorn (in Chinese). China Fruits 2:17–19. Vitamin C Total Soluble Song, W. and S. Lin. 1985. Studies on chromosome (mg/100 g flavonoids sugar Acid numbers of some species of Chinese hawthorn Cultivar fresh fruit) (%) (%) (%) (in Chinese). Acta Hort. Sinica 2:73–76. Da Wang 66.2–83.8 0.41 8.6–10.2 3.13–4.59 Wan, Z. and L. Cai. 1986. Investigation on haw- Ye He 72.9–74.4 0.49 10.7 3.49 thorns of Hubei Province (in Chinese). J. Central Ji An Zi Ro 118.0 0.29 7.4 2.85 China Agr. Univ. 4:400–402. Xi Feng Hong 72.1 0.65 7.5–9.4 1.60–3.20 Wang, M. and P. Yang. 1993. A new variety of Liao Hong 82.1 0.77 10.3 3.56 Crataegus cuneata Siebet Zucc (in Chinese). J. Yan Yang Hong 61.9 12.5 3.34 Northwest For. College 4:96–97. Yan Yang Qing 75.0 0.47 12.0 3.80 Wen, S. 1960. Major hawthorn cultivars of Liaoning Jin Xing 72.9 0.43 10.0 3.65 Province (in Chinese). China Fruits 2:28–30. Han Lu Hong 83.0 0.79 9.4 3.63 Xie, Y. 1984. Analysis of chemical composition of Mian Zha 77.9–87.8 0.78 16.8 3.93 Chinese Crataegus species (in Chinese). Acta Hong Yang Mian 60.4–68.9 0.38 9.8 2.73–3.47 Bot. Sinica 5:383–387. Bai Yang Mian 51.9–69.5 0.45 9.2–9.5 3.56–3.69 Xin, X. and Y. Zhang. 1984. Hawthorn cultivars of Chang Ko 69.3 0.43 8.4 4.04 China and their scientific names (in Chinese). J. Da Jin Xing 62.4–73.6 0.42 4.9 3.57 Shenyang Agr. College 1:76–78. Yu Bei Hong 74.3 0.69 13.8 1.13–2.20 Yen, W. 1982. Species and chemical composition of Yan Guo Hong 70.5 0.64 8.4 3.83 Crataegus in northeastern China (in Chinese). J. Qu Jin Xing 60.9–62.7 0.50 11.3 3.89 Spec. & Valuable Wild Animals & Plants 3:18– Shuang Hong 64.4–68.2 0.45 9.7 2.56 23. Fu Li Hong 24.5–74.3 8.9–9.0 2.70 Yu, D. 1977. Hawthorn, p. 147–164. In: Ji Fu Yi Hao 22.2 5.8 1.16 of fruit crops in China (in Chinese). Agr. Publ. Zuo Fu Yi Hao 34.8 0.22 7.5 1.51 China, Beijing. Zhang, J., B. Feng, and D. Zhang. 1982. Hawthorn (in Chinese). Agr. Publ. China, Beijing. p. Using iosperoxidase analysis to study taxonomic Liu, X. 1990. Germplasm study of wild hawthorn in 11–22. system and relationship of Crataegus pinnatifida Jilin Province (in Chinese). MS Thesis, Jilin Zhang, Y. and X. Xin. 1986. Studies on chromo- (in Chinese). J. Spec. & Valuable Wild Animals Agr. Univ., Changchun. some numbers and isoperoxidase of Chinese & Plants 3:15–19. Nie, S. and B. Ren. 1982. Notes on woody plants of hawthorn (in Chinese). Fruit Crops of Liaoning Guo, T., P. Jiao, J. Li, B. Feng, and X. Sun. 1992. rose family in the Northeast (in Chinese). Bul. 1:16–23. Study of taxonomic system and relationship of Bot. Res. 1:45–51. Zhao, W. 1979. Major hawthorn cultivars of Crataegus pinnatifida (in Chinese). 1992 Excel- Pu, F., S. Lin, and D. Zhang. 1987. Observation on Shandong Province (in Chinese). China Fruits lent Publ. of Hort. Soc. Jilin Province. chromosome numbers of several species and 2:29–31.

1134 HORTSCIENCE, VOL. 30(6), OCTOBER 1995