یادداشت‌هایی بر Aegilops cylindrica (Triticeae, Poaceae) در ایران

نویسندگان

گروه زیست‌شناسی، دانشکده علوم، دانشگاه اصفهان، اصفهان، ایران

چکیده

تحقیق حاضر مبتنی بر مطالعه تاکسونومی 100 نمونه جمعیتی از گونه Aegilops cylindrica جمع‌آوری شده از سرتاسر کشور و همچنین مطالعه نمونه‌های تیپ و نمونه‌های مربوط موجود در هرباریوم‌های موزه تاریخ طبیعی وین و برلین است. نتایج این مطالعه نشان داد که صفات ریخت‌شناختی نظیر: طول سنبله، طول سیخک، تعداد سنبلک‌ها در هر سنبله، حضور/غیبت کُرک روی سنبلک‌ها و محور در بین نمونه‌های جمعیتی مطالعه شده تنوع بالایی دارد. بر اساس نتایج پژوهش حاضر، گونه A. cylindrica در ایران با سه واریته که همگی گزارش جدید برای ایران هستند، شناخته می‌شود.

کلیدواژه‌ها

موضوعات


عنوان مقاله [English]

Notes on Aegilops cylindrica (Triticeae, Poaceae) in Iran

نویسندگان [English]

  • Sayed Zabihollah Hosseini
  • Mohammad Reza Rahiminejad
Department of Biology, Faculty of Sciences, University of Isfahan, Isfahan, Iran
چکیده [English]

The present paper deals with the study of the taxonomy of one hundred accessions of Aegilops cylindica collected all around the country, and also Iranian materials and type specimens and related species housed in the Natural History Museum Vienna (W), the Botanic Garden and Botanical Museum Berlin (B), were studied morphologically. The results showed that morphological traits such as length of spike, length of awn, number of spikelet in each spike, presence/absence of hair on spikelets and rachis were highly variable among the studied materials. Based on the results of this study A. cylindica was recognized as having three varieties, all new records, for Iran.

کلیدواژه‌ها [English]

  • Aegilops cylindrica
  • Triticeae
  • Poaceae
  • Taxonomy
  • Iran

Introduction

Aegilops cylindrica Host, an annual wild grass and almost close relative to Triticum aestivum L. (bread wheat), occurs throughout Mediterranean, Middle East and Asiatic regions (Linc et al., 1999; Slageren, 1994; Karataglis, 1989). This species with a chromosomal formula of 2n = 4x = 28 (DDCC) has been considered as an allotetraploid species resulted from a natural hybridization between A. caudata L. (2n = 2x = 14, CC) and A. tauschii Coss. (2n = 2x = 14, DD) (Linc et al., 1999; Karataglis, 1989). Jaaska (1978) suggested that this species originated from east parts of Turkey, where the distribution areas of its putative parents might had been overlapped.

In his account Tzvelev (1976) recognized four varieties for A. cylindrica: cylindrica, aristulata (Zhuk.) Tzvel., pauciaristata Eig. and prokhanovii Tzvel. occurring in (the former) Soviet union (USSR); a fifth one, i. e., var. rumelica was added by Velenovsky for Flora Bulgarica (Velenovsky 1891). Bor (1970) recognized this species with no infra-specific subdivisions for the Flora Iranicaareas (Iraq, Afghanistan and NW, W of Iran). Literature review shows that the Iranian materials of this species were the subject of taxonomic (Keshavarzi et al., 2006; Arzani et al., 2006; Kharazian, 2008), molecular cytogenetic and C-banding (Linc et al., 1999; Bordbar et al., 2011), isozyme (Jaaska, 1981), and inter-generic hybridization (Morrison et al., 2002) studies. Aegilops cylindrica is also important from the breeding point of view, due to the gene flow between this species and T. aestivum (Guadagnuolo et al., 2001).

This study aims to review the taxonomic status of A. cylindrica in Iran.

Materials and Methods

A selection of 25 representetives (Table 1) belonging to Aegilops cylindrica from 100 accessions collected during the years 1996-2000 all around Iran were studied. The selected accessions included all distribution areas and main phenotypic variations. In order to provide enough plant materials for the morphological and taxonomic studies, the accessions were grown in the research field of the University of Isfahan. Voucher specimens are deposited in the herbarium of the University of Isfahan (HUI). In addition, all the related materials including the Iranian specimens collected for this study, type specimens of A. cylindrica var. cylindrica (W 2284), A. cylindrica var. pauciaristata (W 1426), A. cylindrica var. rumelica Velen. (W 5377) and two parental species i. e. A. tauschii (W 1973-0008961, W 1970-0016311) and A. caudata (W 0000441) housed in the Natural History Museum Vienna (W) and the Botanic Garden and Botanical Museum Berlin (B) were studied.

 

Table 1. List of Iranian Aegilops cylindrica accessions studied taxonomically

Locality

Accession Number

No.

Lorestan: Khorram Abad to Sepidasht, 1900 m, Rahiminejad and Sahebi

Ag. 43

1

Fars: Dasht Arzhan, 2030 m, Rahiminejad and Sahebi

Ag. 59

2

Kurdistan: Saqez to Marivan, 1500 m, Rahiminejad and Sahebi

Ag. 108

3

Azarbaijan: 15 km remain to Ahar from Tabriz, 1550 m, Rahiminejad and Sahebi

Ag. 148

4

Kurdistan: Piranshahr to Sardasht, 1200 m, Rahiminejad and Sahebi

Ag. 173

5

Kurdistan: Boukan to Mahabad, 1490 m, Rahiminejad and Sahebi

Ag. 240

6

Azarbaijan: Urmieh, 1332 m, Rahiminejad and Sahebi

Ag. 262

7

Chaharmahal va Bakhtiari: Lordegan, Deh Soukhteh, 1770 m, Rahiminejad and Sahebi

Ag. 84

8

Kurdistan: Sanandaj to Sonqor, 1620 m, Rahiminejad and Sahebi

Ag. 90

9

Azarbaijan: Ahar to Kalibar, 1430 m, Rahiminejad and Sahebi

Ag. 106

10

Kohgiluyeh va Boyer-Ahmad: Yasouj to Babameydan, 1180 m, Hosseini and Saeedi, (HUI 17985)

Ag. 470

11

Kohgiluyeh va Boyer-Ahmad: Yasouj to Lordegan, Gharah village, 1950 m, Hosseini and Saeedi

Ag. 476

12

Alborz: Karaj to Chalous, Marzan Abad, 500 m, Hosseini and Saeedi

Ag. 478

13

Isfahan: Semirom, 2300 m, Hosseini and Saeedi

Ag. 474

14

Khorasan: Azadshaher to Bojnourd, 1450 m, Hosseini and Saeedi

Ag. 479

15

Khorasan: Shirvan to Mashhad, 1090 m, Hosseini and Saeedi

Ag. 484

16

Khorasan: Neyshabour to Sabzevar, 1080 m, Hosseini and Saeedi

Ag. 488

17

Khorasan: Sabzevar to Shahroud, Mayami, 1120 m, Hosseini and Saeedi

Ag. 489

18

Tehran: Firouzkooh to Tehran, 1810 m, Hosseini and Saeedi

Ag. 490

19

Hamedan: Hamedan to Assad Abad, 2140 m, Rahiminejad and Saeedi

Ag. 497

20

Azarbaijan: Urmieh to Sero, 1286 m, Rahiminejad and Saeedi

Ag. 509

21

Azarbaijan: Ahar to Kalibar, 35 km remain to Kalibar, 1716 m, Rahiminejad and Saeedi, (HUI 17986)

Ag. 516

22

Golestan: Golestan forest, 650 m, Hosseini and Saeedi

Ag. 576

23

Hamedan: Hamedan to Assad Abad, Assad Abad neck, 2140 m, Rahiminejad and Saeedi

Ag. 497

24

Khorasan: Ashkhaneh to Bojnourd, 850 m, Rahiminejad and Saeedi

Ag. 486

25

 

In addition, to certify the ploidy levels of the studied materials, all the accessions were chromosomally examined from the root tips stained in aceto-orcein by squash method (unpublished data). In order to make a comparison among the materials under study, 20 morphological characters (16 quantitative and 4 qualitative) cited in the relevant literatures were evaluted (Table 2).

 

Table 2. Quantitative and qualitative characters considered in study of Aegilops cylindrica varieties in Iran (in quantitative characters means have calculated)

No.

Character

A. cylindrica

var. cylindrica

A. cylindrica

var. prokhanovii

A. cylindrica

var. rumelica

1

Length of spike (cm)

9.36

8.1

5.22

2

Length of spikelet (mm)

10.66

9

10

3

Length of lower most glume (mm)

9

9.1

8.5

4

Width of lower most glume (mm)

3.66

3.4

3.1

5

Length of upper most spikelet awn (cm)

5

4.8

6.63

6

Length of lower most spikelet awn (mm)

2

2.7

3

7

Length of second spikelet awn (mm)

5.33

3

10.7

8

Length of second spikelet glum (mm)

10.33

7

7.2

9

Length of caryopsis (mm)

8.33

6

5.5

10

Width of caryopsis (mm)

2.16

2

2

11

Number of florets in spikelet

3.66

3

3

12

Number of seeds in spikelet

1.66

2

2

13

Number of spikelets in each spike

8

8

5.1

14

Length of second spikelet awn/ Length of second spikelet glum

0.51

0.42

1.48

15

Length of uppermost spikelet awn/ Length of spike

0.53

0.59

1.27

16

Length of lower most spikelet awn/ Length of lower most glume

0.22

0.29

0.35

17

Compact hair on rachis (absent: - / present: +)

-

+

-

18

Hair on glume (absent: - / present: +)

-

+

-

19

Spike shape (oblong: - / cylindric: +)

+

+

-

20

Spike colour (green: -/ brown: +)

-

+

-

 

Results and discussion

The results of this study showed that the geographical distribution of Aegilops cylindrica in Iran was restricted to an area from northwest (Azarbaijan province) eastwardly along Elburz mountains toward northeast (Khorasan province) and southwardly along Zagros mountains toward southwest of Iran with an elevation ranging from 500-2300 m (Figure 1). In comparison with Bor (1970) our observations expanded the distributional range of this species in Iran. Jaask (1978) considered A. cylindrica as a Mediterranean element, where the two putatively diploid progenitor i. e. A. caudata and A. tauschii underwent the hybridization and allopolyploidy processes to create A. cylindrica in Turky; which then emigrated to Europe and Iran.

In A. cylindrica usually the length of awns gradually increased from lowermost spikelet towards the uppermost one in each spike, so that the uppermost spikelet showed the longest awn. One of the studied accessions collected from NW of Iran showed interesting state in which not only was the awn of the uppermost spikelet the longest but also was longer than the total spike’s length. On the other hand, in this specimen all the spikelets in the middle part of the spike showed awns longer than the accessions collected from the other areas. The majority of accessions studied possessed spikes with an average of eight spikelets per spike, however, the least number of spikelets per spike (five spikelets) was observed in above (Table 2).

 

Figure1. A distribution map of Aegilops cylindrica varieties in Iran

 

Also, this study showed that spike shape among the studied materials vary from oblong to cylindrical. All plants having the longest awn and the least spikelets number per spike showed oblong spike shape compared with those having cylindrical spike. These characters were diagnostic features for var. rumelica which we collected from Azarbaijan (Ahar to Kalibar, 35 km to Kalibar, 1716 m) (see Figures 2C, 3, Tables 1, 2).

 

 

Figure 2. Aegilops cylindrica varieties. A. cylindrica (HUI 17989); B. prokhanovii (HUI 17985); C. rumelica (HUI 17986); D. Scabrid surface of glume in var. cylindrica and var. rumelica. E: Densly pilose in var. prokhanovii.

 

Figure 3. A general habit of Aegilops cylindrica Host (var. cylindrica)

 

The presence of scabrid glumes and rachiswas the normal traits in A. cylindrica; neverthless a collected accession from Yasouj to Babameydan (HUI 17985) showed hairy glumes and rachis which is in accordance with Tzvelev (1976) who took the densely short pilose spikelets and rachis variant to describe his new variety (var. prokhanovii) (see Figures 2B, 2D, 2E). Plants of this accession were characterized by brownish spikes and rachis (Table 2).

According to the results of this study, despite the prevalence of hybridization in Poaceae (even inter-generic); from which natural and artificial hybrids between A. cylindrica and T. aestivum are well known and documented (Morrison et al., 2002 and our unpublished data); taxonomic identity of infra-specific taxa of A. cylindrica could be conserved in the sympatric localities; e.g., var. prokhanovii and var. cylindrica grow side by side but still distinct in Yasouj to Babameydan site.

Based on the results of this study we concluded that A. cylindrica grows in Iran with three varieties: cylindrica, prokhanovii and rumelica. A taxonomic key to these varieties was constructed and presented as below:

 

Key to the varieties of A. cylindrica in Iran

1- Awns of the uppermost spikelet longer than the total length of spike; spike usually with 3-5 spikelets …………................................................ var. rumelica
- Awns of the uppermost spikelet shorter than the total length of spike; spike with more spikelets ............................................................................................. 2
2- Spikelets and rachis densely and brownish pilose ................. var. prokhanovii
- Spikelets and rachis scabrid, spikes green ….............................. var. cylindrica

 

Acknowledgments

The authors wish to thank the Office of Graduate Studies of the University of Isfahan for their support. We are also thankful to Dr. Ernst Vitek, Dr. Lia Pignotti, Dr. Robert Vogt and their colleagues in W and B herbaria respectively. Also, we are grateful to Dr. Vello Jaaska, Dr. Bernd R. Frieb and Dr. Van Slageren for their valuable guidance and Ms. Hajian for her precise hand drawing.

References

Arzani, A., Khalighi, M. R., Shiran, B. and Kharazian, N. (2005) Evaluation of diversity in wild relatives of wheat. Czech Journal of Genetics and Plant Breeding 41: 112-117.

Bor, N. L. (1970) Aegilops L. In: Flora Iranica (Ed. Rechinger, K. H.) vol. 70. Akademische Drucku.-Verlagsanstalt, Graz.

Bordbar, F., Rahiminejad, M. R., Saeidi, H. and Blattner, F. R. (2011) Phylogeny and genetic diversity of D-genome species of Aegilops and Triticum (Triticeae, Poaceae) from Iran based on microsatellites, ITS, and trnL-F. Plant Systematics and Evolution 291: 117-131.

Guadagnuolo, R., Savova-Bianchi, D. and Felber, F. (2001) Gene flow from wheat (Triticum aestivum L.) to jointed goatgrass (Aegilops cylindrica Host), as revealed by RAPD and microsatellite markers. Theoretical and Applied Genetics 103:1-8.

Jaaska, V. (1978) Electrophoretic study of phosphatase isoenzyme in the grass genus Aegilopos L. Biochemie und Physiologie der Pflanzen 172: 133-153.

Jaaska, V. (1981) Aspartat aminotransfrase and dehydrogenase isoenzyme: intraspecific differentiation in Aegilops tauschii and the origin of the D genome polyploids in the wheat group. Plant Systematics and Evolution 137: 259-273.

Karataglis, S. S. (1989) Aegilops cylindrica var. kastorianum (Poaceae): A new variety from Greece. Plant Systematics and Evolution 163: 13-20.

Keshavarzi, M., Rahiminejad, M. R. and Kheradmandnia, M. (2007) Discriminative analysis of 10 species of Aegilops L. (Poaceae) native to Iran. Iranian Journal of Science and Technology 31(A3): 321-324.

Kharazian, N. (2007) The taxonomy and variation of leaf anatomical characters in the Genus Aegilops L. (Poaceae) in Iran. Turkish Journal of Botany 31: 1-9.

Kharazian, N. (2008) Chemotaxonomic studies on Aegilops L. (Poaceae) in Iran. Pakistan Journal of Biological Science 11: 1204-1211.

Linc, G., Frieb, B. R. and Kynast, R. G. (1999) Molecular cytogenetic analysis of Aegilops cylindrica Host. Genome 42: 497-503.

Morrison, L. A., Lyzarazu, O. R. and Smith, C. A. M. (2002) Jointed goatgrass (Aegilops cylindrica Host) × wheat (Triticum aestivum L.) hybrid: Hybridization dynamics in Oregon wheat fields. Crop Science 42: 1863-1872.

Slageren, M. V. (1994) Wild wheats. A Monograph of Aegilops L. and Amblyopyrum (Jaub & Spach) Eig (Poaceae). Agricultural university, Wageningen, the Netherlands.

Tzvelev, N. N. (1976) Grasses of the Soviet Union, Part 1. Nauka Publisher, Leningrad, Soviet Union.

Velenovsky, J. (1891) Flora Bulgarica. Prostat Apud Fr. Rivnac, Bibliopolam, Pragae.