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DIVERSITY OF SOIL STREPTOMYCETES AT "EVOLUTION CANYON", MOUNT CARMEL , ISRAEL H. V. VALAGUROVA, V. E. KOZIRITSKAYA, G. A. IUTINSKAYA, A. A. PINDRUS and K. I. ANDREYUK Institute of Microbiology and Virology, Kyiv, Ukraine
Code Number:AC98003 ABSTRACT. Streptomyces complexes of three soil ecotopes at "Evolution Canyon" (Mt.Carmel, Israel) were investigated. Numbers of streptomycetes ranged from 3,000 to 15,000 CFU per g of dry soil. In total eighty-eight species were isolated marked and significant differences between Streptomyces species of the three ecotopes were detected. Streptomycetes are widely distributed in nature. In soil their number varies according to ecological and geographical conditions, soil type, degree of cultivation, water and saline regime, parent rock and mechanical composition. The Streptomyces component of the microbial community increases from northern to southern soils with a corresponding increase of species diversity (Waksman & Starkey, 1931; Waksman, 1959; Krassilnikov, 1970; Andreyuk et al., 1974). In "Evolution Canyon" a combined research program on biodiversity is being carried out, involving a wide range of organisms animals, plants, algae, lichens, mosses and micromycetes (Nevo, 1955; Wasser et al., 1955). The present report deals with the Streptomyces population of the soils of the location. MATERIALS and METHODS Sampling. Evolution Canyon, presumably 3-5 million years old, is located at Lower Nahal Oren, Mount Carmel, Israel and comprises three ecotopes: two slopes, a south-facing slope (SFS) and a north-facing one (NFS) and the valley bottom (VB) with a summer-dry water course. The two slopes are 200 (bottom) and 500 m (top) apart. The SFS is warmer, drier, microclimatically more fluctuating and less predictable than the NFS (Nevo, 1995). The brown-red 'terra rossa' soil of the SFS is covered by thermophilic savannah vegetation while the brown soil of the NFS harbours an oak forest with thermophilic shrubs. The colluvial-alluvial soil of the VB is covered by open forest vegetation. Soil samples were collected at three stations (1, 2 and 3: upper, middle and lower respectively) on the SFS, one at the VB (4) and other three (5, 6 and 7: lower, middle and upper) on the NFS. Samples were collected from the upper layer (0-10 cm) of the soil surface during the dry season (July). Isolation and identification. Samples (10 g) were suspended in sterile distilled water on a reciprocal shaker (120 rpm, 30 min), serially diluted and 0.1 ml of the forth, fifth and sixth dilutions were spread over the surface of mineral medium No. 1 (Gause et al., 1983). Triplicate plates were used for total streptomycete counts after 10 dd incubation at 27 C. Isolates were classified according to the colour of the sporulating aerial and substrate mycelia and transferred to oatmeal agar. Methods suggested by the International Streptomyces Project (Shirling & Gottlieb, 1966) and by Gause et al. (1983) were used for species identification. Species diversity of the isolates was estimated according to Magurran (1991). RESULTS and DISCUSSIONS Streptomycete frequency. The number of streptomycetes ranged from 3,000 to 15,000 CFU per g of dry soil (Table 1) and is slightly lower than in similar soils of Caucasus and Crimea (Andreyuk et al., 1974). The highest values were found in the colluvial-alluvial VB soil. The brown- red SFS soil was poorer than the VB soil but richer than the brown forest NFS soil (an average of 12,900 and 8,100 CFU respectively). The quantity of streptomycetes at three ecotopes is explained by the different type of soils and edaphic conditions (Table 1).
Table 1. Number of streptomycetes (Stm) and some soil characteristics (A.Travlev et al., pers. comm.) of the sampling sites of "Evolution Canyon" (DE: decayed; SD: semidecayed; UN: undecayed).
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Site Stm Humus pH Ammonium Hygro-scopic Plant
x10^3 content, nitrogen, water (%) residues
(%) mg/100g x10^-4
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1 13.3 7.56-6.84 7.83-7.72 0.63-0.68 7.48-6.50 DE & SD
2 11.2 5.49-5.18 7.80-7.30 1.11-0.90 6.47-7.17 DE & SD
3 14.1 6.22-4.87 7.15-7.35 2.75-2.14 6.71-7.16 -
4 15.1 4.55-4.04 7.45-7.65 2.65-3.56 6.46-6.60 DE & SD
5 12.1 11.19-10.15 7.95-7.90 5.34-3.56 7.62-7.59 DE & SD
6 9.1 9.94-9.11 8.15-8.10 5.48-2.84 7.23-7.37 DE, SD & UD
7 3.0 11.81-10.56 7.90 4.74-3.62 - DE, SD & UD
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Streptomycetes develop better under the warm, temperate conditions, at relatively low moisture and neutral or slightly alkaline reaction (Waksman, 1959; Krassilnikov, 1970). Microclimate, nutrient elements, soil reaction and moisture are more favourable for streptomycetes at the VB and SFS sites. The character of vegetation plays a role too. As shown by Waksman & Starkey (1931) streptomycetes numbers are higher in savannah soils than in forest ones. The presence of decayed and semidecayed and absence of non- decayed plant residues at SFS stations are indexes of intensive mineralization processes, possibly by streptomycetes. Species diversity. Eighty-eight Streptomyces species and subspecies were isolated from canyon soils (Table 2). Streptomyces strains with grey aerial mycelium (Cinereus section) were found in all ecotopes and were represented by 45 species. The other sections (Albus, Roseus and Helvolo-Flavus) were represented by 10, 18, and 13 species respectively. Only one species of the Azureus section (Glaucescens series) was isolated at the two canyon slopes, no species of the Coerulescens series of the section were found. Table 2. Streptomyces species isolated at "Evolution Canyon" (Sections and series according to Gause et al., 1983; S.: Streptomyces, A.: Actinomyces; X: found; - : not found)
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Species Station
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1 2 3 4 5 6 7
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SECTION: AZUREUS
Series: Glaucescens
S. glaucus Agre et Preobrazhenskaya 1983 X X X - X X X
SECTION: CINEREUS
Series: Achromogenes
S. aburaviensis Nishimura et al. 1957 X - - - - - -
S. albogriseolus Benedict et al. 1954 - X - - - - -
S. bikiniensis Johnstone et Waksman 1947 - X - - - - -
S. canus Heinemann et al. 1953 - - X X - - -
S. cinereorectus Terechova et Preobrazhenskaya
1983 - - - X - - -
S. clavuligerus Higgens et Kastner 1971 - - - - X - -
S. diastatochromogenes (Krainsky 1914) Waksman
et Henrici 1948 X - - X X - -
S. gardneri (Waksman 1942) Waksman 1961 - - X - - - -
S. gelaticus (Waksman 1923) Waksman et
Henrici 1948 - - - X - - -
S. mutomycini Gause et Maximova 1983 - - - - - X -
S. nodosus Trejo 1961 - - - - X - -
S. pristinaespiralis Mancy et al. 1962 - - - - X - -
S. pseudogriseolus Okami et Umezawa 1955 X X - X - - -
S. spororaveus (Krassilnikov 1970) Gause et al.
1983 - - - X X - -
S. wedmorensis (Waksman 1953) Gause et al. 1983 X - X - - X X
Series: Aureus
S. chattanoogensis Burns et Holtman 1959 - - - X - - -
S. flavovirens (Waksman 1923) Waksman et
Henrici 1948 - - - - X - -
S. griseoflavus (Krainsky 1914) Waksman et
Henrici 1948 - - - - X - -
S. loidensis (Millard et Burr 1926) Waksman 1953 - - - - - - X
S. ostreogriseus Whitefield et al. 1958 X - - - - - -
S. thermoflavus (Kudrina et Maximova 1963)
Pridham 1970 X - X - - - -
S. viridifaciens Pridham et al. 1958 - X - - - - -
Series: Chrysomallus
S. mutabilis (Preobrazhenskaya et Ryabova 1957)
Pridham et al. 1958 - - - X - - -
S. olivaceoviridis (Preobrazhenskaya et Ryabova
1957) Pridham et al. 1958 X - X X X - -
S. viridobrunneus (Krassilnikov 1970) Gause
et al. 1983 - - - X - - -
S. viridogenes (Millard et Burr 1926) Waksman
1953 - - - X X - -
Series: Chromogenes
S. achromogenes Okami et Umezawa 1953 - - - - - X -
S. carpaticus Maximova et Terekhova 1983 - - - - - X -
S. chromofuscus (Preobrazhenskaya et al. 1957)
Pridham et al. 1958 - - - X - - -
S. eurythermus Corbaz et al. 1955 - - X - - - -
S. fumosus (Krassilnikov 1941) Waksman 1953 X - - X - X -
S. misakiensis Nakamura 1961 - - - - - X X
S. nashvillensis McVeigh et Reyes 1961 - - - - - - X
S. roseogriseus Routien 1963 - - - - - X -
S. xanthocidicus Asahi et al. 1966 X - - - - - -
Series: Violaceus
A. albocyaneus Krassilnikov et Agre 1960 - - - - - - X
S. althioticus Yamaguchi et al. 1957 - - X - - - -
S. ciscaucasicus Sveshnikova 1983 - - - X - - -
S. coelicoflavus (Krassilnikov 1970) Terekhova
1983 - - - X - - -
S. galilaeus Ettlinger et al. 1958 - - - - - X X
S. glomeratus Gause et Sveshnikova 1983 - - - X - - -
S. herbaricolor Kawato et Shinobu 1959 - - - X - - -
S. ramulosus Ettlinger et al. 1958 X X - - - - -
S. violaceochromogenes (Ryabova et.
Preobrazhenskaya 1957) Pridham 1970 - - X - - - -
S. violaceorubidus Gause et al. 1983 - - - X - - -
SECTION: ROSEUS
Series: Lavendulae-roseus
S. cattleya Kahan et al. 1979 - - - - - X -
S. filamentosus Okami et Umezawa 1953 - - X - X - -
S. lincolnensis Mason et al. 1963 - - - - X - -
S. roseolilacinus (Preobrazhenskaya et
Sveshnikova 1957) Pridham et al. 1958 - - - X - X X
S. syringini (Preobrazhenskaya et Sveshnikova
1957) Pridham et al. 1958 - - - - X - -
S. toxytricini (Preobrazhenskaya et Sveshnikova
1957) Pridham et al. 1958 - - X - - - -
S. venezuelae Ehrlich et al. 1948 X - - - X X X
Series: Fradiae
A. albilanatus Zhukova et al. 1975 - - - X - - -
S. flavofungini Szabo et Preobrazhenskaya 1983 - - - X - - -
S. fradiae (Waksman et Curtis 1916) Waksman et
Henrici 1948 - - - - - X -
Series: Fuscus
S. alanosinicus Thiemann et Beretta 1966 X - - - - - -
A. chromopurpureus Krassilnikov 1970 - - - - X - -
S. exfoliatus (Waksman et Curtis 1916) Waksman
et Henrici 1948 - - X - - - -
S. moderatus Reusser 1967 - - X - - - -
Series: Roseoviolaceus
A. pallidoviolaceus Sveshnikova et al. 1967 - X - - - - -
S. violaceus (Rossi-Doria 1891) Waksman 1953 - X X - - X -
Series: Ruber
S. aurantiacus (Rossi-Doria 1891) Waksman 1953 - - - X X X -
S. peucetius Grein et al. 1963 - - - - - - X
SECTION: HELVOLO-FLAVUS
Series: Flavus
S. kanamyceticus Okami et Umezawa 1957 - - - - X - -
Series: Helvolus
A. abikoensum subsp. spiralis Kudrina 1957 X - - - - - -
S. albidochromogenes Gause et al. 1983 X - - - - - -
A. cremeospinus Gause et al. 1975 - X - X - X -
S. erythraeus (Waksman 1923) Waksman et Henrici
1948 X - - - - - -
A. flavescens Krassilnikov et al. 1965 - - - - - X X
S. globisporus subsp. tundromycini (Kovalenkova
1957) Pridham 1970 X - - - - - -
S. helvaticus (Krassilnikov et al. 1965) Pridham
1970 - - - X - - -
S. longisporoflavus Waksman 1953 X - X - - - -
S. pluricolorescens Okami et Umezawa 1956 - - - X X - -
S. rubiginosohelvolus (Kudrina 1957) Pridham
et al. 1958 - - - X - - -
A. viridovulgaris Krassilnikov 1970 - X - - - - -
A. viridovulgaris subsp. albomarinus Solovieva
et Fadeeva 1973 X - - - - - -
SECTION: ALBUS
Series: Albus
S. alborubidus (Kudrina 1957) Pridham et al.1958 - - - X - - -
S. albus (Rossi-Doria 1891) Waksman et Henrici
1943 - - - - - X -
S. candidus (Krassilnikov 1970) Gause et al.1983 - - - - - X X
S. longisporus (Krassilnikov 1941) Waksman 1953 - - - - X - -
S. sindenensis Nakazawa et Fujii 1957 - - - - X X -
Series: Albocoloratus
S. coralus Dietz 1964 X - - X - - -
S. griseoloalbus (Kudrina 1957) Pridham et al.
1958 X - - X X X -
A. imbricatus Tsyganov et al. 1970 X X - X X - -
S. monomycini Gause et Terekhova 1983 X - - - - - -
S. rimosus Sobin et al. 1953 - - - X - - -
SECTION: IMPERFECTUS
S. vendargus Pridham et al. 1958 X - - - - - -
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In regard to distribution inside the system, in the different ecotopes the following generalisations are possible. SFS soil is characterized by the absence of species of the Albus, Fradiae, Ruber and Flavus series of species. Fuscus, Roseoviolaceus, Flavus and Glaucescens series species are also absent in VB soils. Species of all sections could be isolated from NFS soils. Forty-two species were isolated from NFS, 40 from SFS and 31 from VB soils. A similar relationship had previously been noted for other cryptogamic organisms (Wasser et al., 1955). In detail only 7 species are common to all studied ecotopes, 15 are found only in the colluvial-alluvial soil, 26 are present in brown forest soils and 25 species seem specific to the brown-red soils. These data, stress a low similarity of the Streptomyces population in the three ecotopes. Distribution of the species according to their frequency (Fig. 1) shows that in SFS soil there is one species (S. violaceus) with relatively high abundance and a group of organisms of lower frequency. Curves of VB and NFS Streptomyces complexes are similar, more compact in distribution and characterized by the absence of dominating species.
Differences between the population composition at the three ecotopes are confirmed by other parameters (Magurran, 1991), such as Margalef's index (7.0, 7.2 and 8.5 for SFS, VB and NFS respectively), Berger-Parker's diversity index (8.77, 12.99 and 14.28) and Simpson's diversity index (2.38, 3.22; and 3.85). On the other hand, Jakkard's and Sorensen's similarity indexes, are rather low (0.09 to 0.32). In conclusion streptomycetes are components of the microbial cenosis of all three soil types. At the same time they show significant quantitative and qualitative differences as shown by other organisms (Wasser et. al., 1995; Nevo, 1955). ACKNOWLEDGEMENTS. We thank prof. E. Nevo, Institute of Evolution, University of Haifa, Israel, for the soil samples and the Director of the Institute of Biology of Dnepropetrovsk State University, prof. A. Travleev, and his collaborators for soil analyses. REFERENCES
Andreyuk, K. I., H. V. Vladimirova & S. B. Kogan (1974). Soil Actinomycetes of South European Part of USSR and their Biological Activity. Naukova Dumka Publ., Kyiv (in Russian) Gause, G. F., T. P. Preobrazhenskaya, M. A.Sveshnikova, L. P. Terekhova & T. S. Maksimova (1983). Key to Actinomycetes. Nauka Publ., Moscow. (in Russian) Krassilnikov, N. A. (1970). The Actinomycetes. Nauka Publ., Moscow (in Russian) Magurran, A. E. (1991). Ecological Diversity and its Measurement. Chapman & Hall, London Nevo, E. (1995). Asian, African and European biota meet at 'Evolution Canyon' Israel: local tests of global biodiversity and genetic diversity patterns. Proc. R. Soc. Lond. B, 262: 149-155 Shirling, E. B. & D. Gottlieb (1966). Methods for characterization of Streptomyces species. Intern. System. Bacteriol., 16: 313-340 Waksman, S. A. (1959) The Actinomycetes. Nature, Occurrence, and Activities. The Williams & Wilkins Co., Baltimore Waksman, S. A. & R. L. Starkey (1931). The Soil and Microbe. J.Wiley & Sons, New York Wasser, S. P., E. Nevo, O. N. Vinogradova, I. L. Navrotskaya, I. A. Ellanskaya, P. A. Volz, V. M. Virchenko, P. M. Tsarenko & S. Ya. Kondratyuk (1995). Diversity of cryptogamic plants and fungi in 'Evolution Canyon', Nahal Oren, Mount Carmel Natural Preserve, Israel. Isr. J. Plants Sc., 43: 367-383 Copyright 1998 C.E.T.A., The International Centre for Theoretical and Applied Ecology, Gorizia The following images related to this document are available:Line drawing images[ac98003a.gif] |
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