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Laboratory of Forest Ecology, Graduate School of Agriculture, Kyoto University, Kyoto 606-8502 Japan
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ABSTRACT |
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Decomposition processes of Swida controversa leaves were investigated in initially sterilized (fungi-excluded) and nonsterilized freshly fallen leaves to examine the relationship between chemical changes and fungal succession during decomposition and the effect of exclusion of previously established phyllosphere fungi from leaves on subsequent decomposition and fungal succession. Fifteen species were isolated frequently from decomposing leaves with surface-disinfection and washing methods. These fungi were divided into early and late colonizers according to their occurrence during decomposition. The 1.5 y decomposition process was divided into three stages characterized by different dominant organic chemical constituents. A clear relationship was demonstrated between chemical changes and fungal succession. Total hyphal length and frequencies of some early colonizers were reduced in initially sterilized leaves at 3 wk, but this had no significant effect on loss of litter mass or chemical changes during the first 3 wk or on the subsequent decomposition and fungal succession.
Key words: dogwood, hyphal length, lignin, litter bag, phyllosphere
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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, Kjøller and Struwe 1982, Cooke and Rayner 1984). Fungal succession during decomposition has been well documented in terms of total mycelial abundance (Berg 1991, Osono and Takeda 2001b, Osono et al 2003b) and species composition (Hudson 1968). Few studies, however, have examined functional relationships between fungal succession and changes in carbon fractions and major nutrients such as nitrogen during decomposition (Osono and Takeda 2001b). Further studies are necessary to follow the chemical and biological aspects of decomposition simultaneously and examine the possible role of fungal succession in litter decomposition processes.
Chemical decomposition of leaf litter follows a sequential pattern with different classes of organic compounds dominating the process as it proceeds (Berg and McClaugherty 2003; Osono and Takeda 2005a, b). In general loss of soluble components occurs in the first stage, followed by holocellulose decomposition in the second stage. Finally lignin becomes a dominant component in the third stage when litter mass loss slows and litter approaches humus.
Studies of fungal succession have emphasized the persistence of endophytic and epiphytic phyllosphere fungi from live leaves to freshly fallen leaves (Hudson 1968) and their frequent occurrence in early stages of decomposition when litter mass loss and chemical changes take place most rapidly (Osono 2002, Osono et al 2004, Koide et al 2005a, b). Being the first colonizers phyllosphere fungi have the advantage of gaining access to readily available organic compounds in freshly fallen leaves, before fungi that colonize after litter fall (Stone 1987). The sterilization of leaves and exclusion of phyllosphere fungi reduced the decomposition rate of the leaves (Tanner 1981) and might alter substrate use by succeeding fungal decomposers (Osono 2003, Koide et al 2005a), suggesting the potential importance of prior colonization by phyllosphere fungi in initial and subsequent stages of decomposition and fungal succession.
In the present study I investigated decomposition of freshly fallen leaves of giant dogwood (Swida controversa (Hemsley) Sojak) (Cornaceae) with reference to chemical changes and fungal succession in initially sterilized (phyllosphere fungi-excluded) and nonsterilized (control) leaves. The objectives were to examine (i) the relationship between chemical changes and fungal succession during decomposition and (ii) the effect of exclusion of phyllosphere fungi on leaf decomposition. Freshly fallen leaves were sterilized by exposure to ethylene oxide gas so that phyllosphere fungi were excluded without changing the chemical properties of leaves. Decomposition of initially sterilized leaves was compared with nonsterilized leaves to quantify the contribution of phyllosphere fungi to decomposition. Preliminary studies indicated that leaf litter of S. controversa was characterized by low initial lignin content and rapid mass loss and frequently was colonized by phyllosphere fungi (Osono and Mori 2004; Osono et al 2004; Osono and Takeda 2005a, b). Thus I hypothesized that the exclusion of phyllosphere fungi would retard mass loss of initially sterilized leaves compared to nonsterilized leaves.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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. Three study plots of 5 x 5 m2 were laid out in the study site. Each plot was divided into five subplots of 5 x 1 m2 for the decomposition experiment. The three plots were chosen randomly and located within 200 m of each other.
Litter bag method.— –
Decomposition of dogwood leaf litter was studied with a litter bag method (Crossley and Hoglund 1962). Freshly fallen leaves were collected Nov 2001 from forest floor. Leaves were air-dried at room temperature (ca. 15–20 C) 1 wk. The litter (3 g) was enclosed in a litter bag (15 x 15 cm2) made of polypropylene shade cloth with a mesh size of ca. 2 mm. A total of 228 bags were prepared. One-half (114) of these bags was sterilized by exposure to ethylene oxide gas at 60 C for 3 h (denoted as initially sterilized leaves) and the other 114 bags were not exposed to ethylene oxide (denoted as initially nonsterilized leaves). Nine bags with initially sterilized leaves and nine with initially nonsterilized leaves were used for the determination of oven-dry mass at 40 C and initial chemical composition and the other 210 bags for decomposition experiments.
I confirmed that initial contents of lignin, holocellulose, soluble carbohydrates and nitrogen were not significantly different between initially sterilized and nonsterilized leaves (TABLE I; t-test, P > 0.05). Methods for chemical analyses are described below. Before the experiments, initially sterilized leaves (1.0 g) were placed on 2% malt-extracted agar in three Petri dishes (9 cm diam), and after 8 wk incubation at 20 C in the dark no microbial colonies had developed. Thus the effectiveness of the sterilization was confirmed.
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Foreign plant remains attached to the outside of the bags were carefully removed with forceps. Eighteen (nine initially sterilized and nine nonsterilized) of the 30 bags were used for mass determination and chemical analyses. The other 12 (six initially sterilized and six nonsterilized) were used for hyphal length estimation and fungal isolation.
Chemical analyses.— – Remaining mass of litter was determined after drying samples to a constant weight at 40 C, and mean values of remaining mass were calculated for each sampling. The samples were combined for each plot, ground in a laboratory mill to pass a 0.5 mm screen and used for chemical analyses. Analyses were carried out in Jun 2003.
The amount of lignin in the samples was estimated by gravimetry according to a standardized method using hot sulfuric acid digestion (King and Heath 1967). Total carbohydrate content was estimated by the phenol-sulfuric acid method (Dubois et al 1956). Soluble carbohydrate was extracted from the sample with 50% methanol (v/v) at 75 C for 60 min. Soluble carbohydrate content was estimated by the phenol-sulfuric acid method. The holocellulose fraction was calculated as the difference between the total carbohydrate and the soluble carbohydrate. Total N content was measured by automatic gas chromatography (NC analyzer SUMIGRAPH NC-900, Sumitomo Chemical Co., Osaka, Japan). Methods are described in Osono et al (2003a).
Hyphal length estimation.— –
Hyphal length in decomposing leaves was measured with the agar film method of Jones and Mollison (1948) but with several modifications (Osono et al 2003b). The samples were processed within 24 h of sampling. Decomposing leaves from two bags collected from each plot were combined, and 1 g (fresh weight) was homogenized in a blender at 10 000 rev/min in 49 mL of distilled water for 3 min. The suspension (20 mL) was diluted with 20 mL of molten agar solution (final concentration 1.5% [w/v]) and mixed slowly on a magnetic stirring plate. Three agar films were prepared from each suspension in a haemocytometer (0.1 mm depth), transferred to glass slides and dried 24 h. The films were dual-stained with fluorescent brightener (FB) and acridine orange (AO), each for 1 h. Fluorescent brightener binds to chitin in fungal cell walls (West 1988) and enables viewing of all hyaline hyphae that are live or ghost (empty). Acridine orange binds to nucleic acids in live fungal cells (Rost 1992) to view live hyphae.
The stained films were mounted between slides and cover slips with one drop of immersion oil (type DF, Cargille Laboratories Inc., Cedar Grove, New Jersey) and examined with a Nikon Microphot-SA epifluorescent microscope equipped with a high-intensity mercury light source. A Nikon UV-1A filter cube was used for examination of FB-stained hyphae, and a Nikon B-2H filter cube was used for AO-stained hyphae. Darkly pigmented hyphae that were not stained with FB were observed by bright field microscopy. Microscope fields were selected randomly, and 25 fields were observed for each slide at 1000x magnification. Hyphal lengths were estimated with an eye-piece grid and the grid-intersection method (Olson 1950). Total hyphal length was calculated as the sum of FB-stained hyphal length and the darkly pigmented hyphal length. Acridine orange-stained hyphae were regarded as live. Hyphae with clamp connections were classified as Basidiomycota, but it was difficult to estimate hyphal length of individual basidiomycete species in decomposing leaves. In the present study, therefore, length of clamp-bearing hyphae in leaves was estimated as total biomass of basidiomycetes in spite of the fact that this might have resulted in underestimation of basidiomycete biomass because the frequency of clamp formations varies between species. The remaining samples were used for fungal isolation or dried at 40 C for 1 wk to determine water content and convert fresh mass to dry mass. Water content was calculated according to this equation: water content (%) = mass of water (g)/mass of litter (g) x 100.
Fungal isolation.— –
For the isolation of fungi from decomposing leaves, a surface disinfection method (Kinkel and Andrews 1988, Hata 1997) and a modified washing method (Harley and Waid 1955) were used according to Osono (2002). Fungal isolation was carried out within 6 h of sampling. Ten pieces of decomposing leaves (approximately 5 x 5 mm2) were taken from each bag, and five of them were used for the surface disinfection method and the other five for the washing method. Thus a total of 120 pieces (60 from initially sterilized and 60 from nonsterilized leaves) were used for two isolation methods on each sampling.
For surface disinfection, leaf pieces were submerged in 70% ethanol (v/v) 1 min to wet the surface, then surface-disinfected 15 s in a solution of 15% hydrogen peroxide (v/v) and submerged 1 min in 70% ethanol. The pieces were rinsed with sterile, distilled water, transferred to sterile filter paper in Petri dishes (9 cm diam) and dried 24 h to suppress vigorous bacterial growth after plating (Widden and Parkinson 1973). The pieces were placed on 9 cm Petri dishes containing LCA (Miura and Kudo 1970), one piece per plate. LCA contains glucose 0.1%, KH2PO4 0.1%, MgSO4·7H2O 0.02%, KCl 0.02%, NaNO3 0.2%, yeast extract 0.02%, and agar 1.3% (w/v). LCA was used because its low glucose content suppresses overgrowth of fast-growing species and because LCA induces sporulation and is useful for identification (Osono and Takeda 1999).
For modified washing, leaf pieces were washed in a sterile test tube and agitated in a vertical shaker 1.5 min to isolate fungi growing actively on the surface. The pieces were washed serially in five changes of 0.005% Aerosol-OT (Di-2-etylhexyl sodium sulfosuccinate) solution (w/v) and rinsed with sterile distilled water five times. The washed pieces were treated in the same manner as that used in the plating-out procedure of the surface disinfected leaves.
Plates were incubated at 20 C in the dark and observed at 3 d and at 2, 4 and 8 wk after surface disinfection or washing (Osono and Takeda 1999). Any hyphae or spores on the plates were subcultured on fresh LCA plates, incubated and identified.
Data analyses.— – Frequency of single species was calculated as a percentage of the number of leaf pieces with the species out of the 10 pieces tested on initially sterilized or non-sterilized leaves, which were surface-disinfected or washed after harvest in each plot on each sampling. When the frequency of a species was significantly greater (P < 0.05) than 10% by Fisher’s exact probability test, the species was regarded as frequent.
A paired t-test was used to compare mean values of decay constant and hyphal length and frequency of fungi at the first sampling between initially sterilized and nonsterilized leaves. A two-way analysis of variance was performed for frequently isolated fungi to evaluate differences in frequencies of fungi during the study using initial leaf sterilization (initially sterilized and nonsterilized leaves) and isolation method (surface disinfection and washing) as independent variables.
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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). Mass loss was most rapid during the first 3 wk with loss of about 20% of original mass (FIG. 1). The mass of leaves then continued to decrease until 11 mo, and the decrease was gradual from 11 to 18 mo. About 10% of the original litter mass remained after 18 mo. The decay constants as Olson’s k (Olson 1963) were not significantly (paired t-test, P > 0.05) different between initially sterilized leaves (2.03 ± 0.22, mean ± SE, n = 3) and nonsterilized leaves (2.04 ± 0.16, n = 3).
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). Stage I corresponded to the first 3 wk, which was characterized by the rapid decrease of holocellulose, soluble carbohydrate and nitrogen with imperceptible mass loss of lignin. Stage II corresponded to the period between 3 wk and 11 mo, which was characterized by decrease in lignin, holocellulose and nitrogen mass and simultaneous increase in relative lignin and nitrogen content. Stage III corresponded to 11–18 mo, which was characterized by the slow decrease of lignin and holocellulose. These patterns were similar between initially sterilized and nonsterilized leaves (FIG. 2).
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). After 5 mo total hyphal length gradually decreased for both initially sterilized and nonsterilized leaves. Live hyphal length and the percentage live length of total hyphal length were not significantly different (paired t-test, P > 0.05) at 3 wk between initially sterilized and nonsterilized leaves and decreased during decomposition in both initially sterilized and non-sterilized leaves (FIG. 3). Dark-pigmented hyphal length at 3 wk was significantly less (paired t-test, P < 0.05) on initially sterilized leaves than on initially nonsterilized leaves (FIG. 3). Dark-pigmented hyphal length was variable, and the percentage dark-pigmented hyphal length of total hyphal length increased during decomposition in both initially sterilized and nonsterilized leaves. The mean length of hyphae with clamp connections was 112 m/g dry litter (range 0–776 m/g) in both initially sterilized and nonsterilized leaves, comprising a mean of 1.8% (range 0–15.8%) of total hyphal length.
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). For washed decomposing leaves, the number of species increased during the first 9 mo and decreased thereafter (FIG. 4). These patterns were similar between initially sterilized and nonsterilized leaves.
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). These species were divided into early and late colonizers according to their occurrence under the various treatments (surface-disinfected or washed decomposing leaves, which initially were sterilized or nonsterilized) (FIG. 5).
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). The effect of initial leaf sterilization was detected on Phoma sp.1 and Pestalotiopsis sp.2, which were significantly less frequent at 3 wk and (or) over decomposition period on initially sterilized leaves than on initially nonsterilized leaves (TABLE II). Frequencies of three early colonizers were significantly greater on washed decomposing leaves than on surface-disinfected decomposing leaves (TABLE II), but Phoma sp.1 and Cladosporium cladosporioides were still frequent on surface-disinfected decomposing leaves (FIG. 5).
The frequencies of the other 12 fungi increased in 5–12 mo and decreased thereafter or increased as the decomposition progressed and thus were regarded as late colonizers (FIG. 5). Their frequencies were not significantly different between initially sterilized and nonsterilized leaves at 3 wk or during decomposition period (TABLE II). The effect of isolation method was not detected on Arthrinium sp. Frequency of Geniculosporium sp.1 was significantly greater on surface-disinfected decomposing leaves than on washed decomposing leaves. Frequencies of the other 10 species were significantly greater on washed decomposing leaves than on surface-disinfected decomposing leaves (TABLE II), but Trichoderma viride was still frequent on surface-disinfected decomposing leaves (FIG. 5).
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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, Hasegawa and Takeda 1996, Osono and Takeda 2001b, Osono et al 2003b), whereas the decreasing trend of the percentage live length was consistent with these reports. The result that the number of species on washed decomposing leaves was greatest in the middle of the study period appears to reflect the ability of a relatively large number of species to colonize leaves, which is represented by the overlap in the distribution of species in early and late colonizers.
Fungal succession on decomposing Swida leaves followed a pattern described by Hudson (1968). An early colonizer C. cladosporioides was regarded as primary saprophytes according to Hudson (1968), whereas later colonizers such as Trichoderma hamatum, Mucor hiemalis, and Absidia glauca might correspond to secondary sugar fungi because they were frequent in Stage III when lignin was the dominant constituent of decomposing leaves (Hudson 1968, Osono and Takeda 2001b). The hyphae of basidiomycetes, major lignocellulose decomposers of leaf litter (Lindeberg 1946, Miyamoto et al 2000, Osono and Takeda 2002, Osono et al 2003a) accounted for about 2% of total fungal biomass in Swida leaves. This low abundance probably is attributable to low lignin content and high content of nonlignified carbohydrates in leaves that favor the colonization of fast-growing species that excluded slower-growing basidiomycetes. In contrast mycelia of basidiomycetes were abundant in lignin-rich Fagus leaves, accounting for about 20% of total fungal biomass (Osono and Takeda 2001b).
Relationship between decomposition and fungal succession.— –
The present study demonstrates a clear relationship between chemical changes and fungal succession. The three-stage pattern of organic chemical changes has been reported in several litter types (Berg and McClaugherty 2003; Osono and Takeda 2005a, b). The duration of Stage II in Swida leaves was shorter than those of other litter types, probably due to the faster loss of nonlignified holocellulose in Swida leaves (Osono and Takeda 2005a, b).
The fungal colonization accounted for a part of the rapid loss of soluble carbohydrates and nonlignified holocellulose in Stage I because the highest abundance of live mycelium was recorded in this stage, reflecting the highest potential of fungi to decompose these carbohydrates (Ingham and Klein 1984). The early colonizers might be responsible for hyphal length and decomposition in Stage I. In fact species in Cladosporium have shown to use highly available energy sources such as soluble and nonlignified carbohydrates (Hudson 1968, Osono and Takeda 2002, Osono et al 2003a).
The constant mass loss of both holocellulose and lignin during Stage II indicates that fungi might have actively used these components. Cellulolytic Trichoderma spp. probably were responsible for mass loss of nonlignified holocellulose. Geniculosporium sp. (Xylariaceae) likely was responsible for the mass loss of lignocellulose (Hering 1967, Hudson 1968, Osono and Takeda 2002). Low lignin content in Swida leaves might enhance lignocellulose decomposition by this xylariaceous fungus (Osono and Takeda 2001a). The relative increase of lignin content during Stage II indicated that the fungal community preferred holocellulose to lignin.
Effect of exclusion of phyllosphere fungi.— –
The initial reduction of fungal colonization in initially sterilized leaves had no significant effects on litter mass loss and chemical changes in Stage I. Litter mass loss in Stage I was not only accounted for by fungal activity but likely also by physical leaching of soluble substances (Tietema and Wessel 1994, Berg and McClaugherty 2003). The relatively high initial content of soluble carbohydrates in Swida leaves suggests that nonbiological leaching accounted for the dominant part of the mass loss during Stage I and that the contribution of fungal activity to the mass loss was smaller than leaching even in initially nonsterilized leaves. Initial leaf sterilization also had negligible effect on the subsequent decomposition and fungal succession in Stages II and III. This probably is attributable to the result that the decomposition of holocellulose and lignin was carried out by late colonizers that replaced early colonizers when the highly available carbohydrates were exhausted by leaching and fungal activity. In summary the initial sterilization of leaves and exclusion of previously establishing phyllosphere fungi resulted in the reduction of initial colonization of some fungi in decomposing leaves but such reduction had negligible effects on subsequent decomposition and fungal succession on S. controversa leaf litter.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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1 Corresponding author. E-mail: fujijun{at}kais.kyoto-u.ac.jp
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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initially nonsterilized leaves.
