보고서 정보
주관연구기관 |
고려대학교 Korea University |
연구책임자 |
김진수
|
보고서유형 | 최종보고서 |
발행국가 | 대한민국 |
언어 |
한국어
|
발행년월 | 2012-03 |
과제시작연도 |
2011 |
주관부처 |
산림청 |
사업 관리 기관 |
산림청 Korea Forest Service |
등록번호 |
TRKO201400015530 |
과제고유번호 |
1405001294 |
DB 구축일자 |
2014-09-20
|
키워드 |
산림식물.유전자원보존.표본추출전략.유전다양성.생태적 특성.Forest plant.Genetic conservation.Sampling strategy.Genetic diversity.Ecological characteristics.
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초록
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○ 산림식물 보존전략 수립을 위한 분포특성 조사
- 54개 분류군 (목본 15종, 초본 39종)에 대한 분포특성 조사
- 종별 자생지에서 고도, 방위 등을 포함하는 환경특성, 종다양도, 우점도 등을 포함하는 식생특성, 토성, 유기물함량, 유효인산 등을 포함하는 토양 특성 조사
○ 유전다양성에 근거한 산림식물 표본추출 전략 개발
- 유전다양성 자료를 이용하여 개체 또는 집단 추출을 통해 확보되는 유전다양성 정도를 시뮬레이션 할 수 있는 컴퓨터 프로그램 SGD (Sampling Genetic Diversity) 개
○ 산림식물 보존전략 수립을 위한 분포특성 조사
- 54개 분류군 (목본 15종, 초본 39종)에 대한 분포특성 조사
- 종별 자생지에서 고도, 방위 등을 포함하는 환경특성, 종다양도, 우점도 등을 포함하는 식생특성, 토성, 유기물함량, 유효인산 등을 포함하는 토양 특성 조사
○ 유전다양성에 근거한 산림식물 표본추출 전략 개발
- 유전다양성 자료를 이용하여 개체 또는 집단 추출을 통해 확보되는 유전다양성 정도를 시뮬레이션 할 수 있는 컴퓨터 프로그램 SGD (Sampling Genetic Diversity) 개발
- ISSR 및 동위효소 표지자를 이용하여 22개 분류군 (목본 7종, 초본 15종)의 유전분석을 수행하고 보존을 위한 기본 자료 제공
- 유전다양성 및 생태적 특성에 근거하여 22개 모델종의 보존을 위한 적정 표본크기 추정
- 연구결과를 토대로 산림식물 유전자원 보존을 위한 표본추출 기본전략을 적정 집단 수 및 개체수를 중심으로 제시
Abstract
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1. Investigation of ecological characteristics of 54-model species
This study intended to understand the ecological characteristics of the 54 taxa, including 15 taxa of woody plants and 39 taxa of herbaceous plants (Table 1). The 54 taxa include 10 Korean endemic plants and 34 rare plants: criti
1. Investigation of ecological characteristics of 54-model species
This study intended to understand the ecological characteristics of the 54 taxa, including 15 taxa of woody plants and 39 taxa of herbaceous plants (Table 1). The 54 taxa include 10 Korean endemic plants and 34 rare plants: critically endangered (CR, 3), endangered (EN, 8), vulnerable (VU, 11), near threatened (NT, 11), and data deficient (DD, 1).
To identify the ecological characteristics of target species, basic data (i.e., altitude, slope degrees, slope direction, litter depth, and illuminance) were investigated by habitat. In addition, to identify the vegetation of habitats, the species richness, species diversity, dominance, and evenness were estimated using coverage and frequency of total species. The soil characteristics were measured by soil texture, field capacity, organic matter, pH, available phosphorus, and salinity.
Environmental characteristics were summarized as altitude, slope direction, and relative illuminance. First, 15 of 54 taxa grew at the area 600 m or higher; 19 taxa at the area less than 600 m high; and the remaining 20 taxa at the area of altitude ranging 100–1000 m with vertical distribution (Table 1). With the slope directions of habitats in each taxon, excluding Nymphoides peltata, 28 taxa were mainly located on the northern to western slope, 12 taxa were located on the eastern to southern slope, and the remaining 13 taxa were located in all directions (Table 1). For 12 woody plants, the relative illumination of the habitats was investigated. Three taxa of the 12 woody plants (Daphne kiusiana, I lex crenata, and Rhododendron brachycarpum) were found in the area with 50% or less relative illumination, where shade was maintained by overstory vegetation. Three other taxa (Betula ermanii, Syringa wolfii, Berberis amurensis var. quelpaertensis) were distributed in the area with 90–100% relative illumination, where shade was very little. The remaining 6 taxa (Exochorda serratifolia, Lonicera vesicaria, Zanthoxylum planispinum, Aristolochia manshuriensis, Forsythia ovata, Vitex negundo var. incisa) were found in areas with varying illumination.
Vegetation analysis showed that dominant species in overstory layer were varied by species. Nine taxa (Megaleranthis saniculifolia, Symplocarpus nipponicus, Hanabusaya asiatica, Corydalis grandicalyx, Epimedium koreanum, Iris odaesanensis, Lilium distichum, Viola diamantiaca, and Saussurea calcicola) were usually distributed in Quercus mongolica forest. Five taxa (Aristolochia manshuriensis, Viola websteri, Eleutherococcus senticosus, Astilboides tabularis, and Scopolia japonica) were usually discovered in Carpinus cordata, Juglans mandshurica, and Acer pictum subsp. mono forest. Meanwhile, 12 taxa (Syringa wolfii, Berberis amurensis var. quelpaertensis, Vitex negundo var. incisa, Wedelia prostrata, Nymphoides peltata, Crypsinus hastatus, Patrinia rupestris, Bupleurum euphorbioides, Pyrrosia petiolosa, Aeginetia indica, Fallopia sachalinensis and Sarcanthus scolopendrifolius) showed the lack or very low level of coverage in tree layer. The other taxa showed various vegetation structures by species and habitat environment (limestone region, island, etc.). Species diversity indices were higher for forest environment than for coastal sand or aquatic ecosystem. In addition, the taxon that grew on various habitat slopes without preference for direction (i.e., northern-western direction or southern-eastern direction) showed higher species diversity than others. For most species, the species dominance was less than 0.3 and evenness was higher than 0.8.
For soil characteristics, the organic matter, field capacity, pH, soil texture, and available phosphorus were investigated. The organic matter showed positive correlation with field capacity and these two values were higher for areas near the valley than for those near the mountain ridge. Soil acidity (pH) was distinctive among taxa by parent material of soil (acidic soil region versus limestone region).
Soil types were mostly sandy loam, loam, and slit loam. Available phosphorus was greatly different between species and populations; no distinctive correlation was found between the available phosphorus and other factors. For some of the taxa (Syringa wolfii, Lonicera vesicaria, and Berberis amurensis var. quelpaertensis) that were distributed mostly in high mountains, distinctive soil characteristics were found. In this area, clay content was low, organic matter content was high, and acidification of soil was found in progress.
As such, habitat environment and ecological characteristics of 54 taxa were identified. The data on 22 taxa for which genetic analysis was performed were used as basic data for developing a sampling strategy and for preserving genetic resource. Besides, all these results are expected to be useful for research and the preservation of relevant species.
2. Development of sampling strategy for genetic conservation of forest plant
2.1 SGD - A simulation program for sampling genetic diversity
To test the optimum sample size, the simulation program, Sampling Genetic Diversity (SGD), which enables the sequential sampling of the desired number of populations or individuals, was developed. Using this program, the changes to the extent of genetic diversity, which is secured by increasing certain number of populations or individuals in a sample, can be estimated. According to as many iterations of random sampling as desired, the SGD program shows not only the mean values but also the ranges of diverse parameters of the genetic variation included in each sample. Finally, the efficiency of the sampling is evaluated based on the relative amount of genetic variation secured in each sample to that of the total amount of genetic variation revealed by the allozyme and ISSR markers in each species.
2.2 Genetic diversity and structure of 22-model species
This study was conducted to figure out the genetic diversity and structure in 22-model species (7 woody species and 15 perennial herbs) selected by considering mainly their life history, breeding systems, and distribution patterns. The 22-model species were analyzed by ISSR marker and 11 of 22 species were analyzed by allozyme marker, additionally. For several model species, especially those propagated vegetatively, the clonal diversity and spatial genetic structure (SGS) were also investigated. This information on the genetic diversity and structure in each species was considered the ultimate goal to be achieved by employing different sampling methods for genetic conservation.
Based on the ISSR marker, percent of polymorphic loci at the population level varied from 27.8% (Echinosophora koreensis) to 82.5% (Abelia tyaihyoni ) with an average of 61.5% among 7 woody plant species. The gene diversity (hp) ranged from 0.101 (Echinosophora koreensis) to 0.285 (Syringa wolfi ) with an average of 0.189.
Percent of polymorphic loci was variable among 15 herbaceous plants ranging from 5.5% (Viola websteri ) to 90.2% (Lilium distichum) with average of 60.1%. Gene diversity was varid from 0.013 (Viola websteri ) to 0.279 (Corydalis grandicalyx) with an average of 0.203. Compared to the previous results of RAPD analysis from across many plant species (Nybom, 2004), the genetic diversity obtained from the present study was comparable with them or slightly lower.
In the case of allozyme marker, percent of polymorphic loci at the population level showed a range from 2.1% (Viola websteri ) to 78.0% (Lilium distichum) with an average of 53.1% among 9 herbaceous plants. Average number of alleles per locus ranged from 1.02 (Viola websteri ) to 2.52 (Lilium distichum) with an average of 1.81.
Expected heterozygosity was also varied among species ranging from 0.006 (Viola websteri ) to 0.228 (Symplocarpus nipponicus) with an average of 0.138. Compared to the previous study on Korean native plants, the genetic diversity measured in this study among 9 species (Hep=0.138) was very similar with them (Hep=0.139). Genetic diversity in Korean native plant species estimated in the present study and previous studies were significantly higher compared with those of other species abroad (Hamrick and Godt, 1990).
Of the 11 species for which analysis was performed with allozyme marker, 9 species excluding Megaleranthis saniculifolia and Viola websteri had mean FIS value of–0.009 (Hanabusaya asiatica) to 0.094 (Symplocarpus nipponicus), almost approximating to the Hardy-Weinberg equilibrium. Megaleranthis saniculifolia and Viola websteri showed homozygote excess with FIS value of 0.258 and 0.523, respectively. The two species were considered to be greatly influenced by selfing or inbreeding.
In comparison of interpopulational differentiation according to the model species, the ФST values based on the ISSR marker ranged from 0.068 (Betula ermanii ) to 0.749 (Daphne kiusiana) with an average of 0.280 among 7 woody plant species. In the case of 15 herbaceous species, the genetic differentiation (ФST) varied between 0.054 (Corydalis grandicalyx) and 0.908 (Viola websteri ) with an average of 0.302.
Based on the allozyme marker, FST values varied between 0.022 (Lilium distichum) and 0.851 (Viola websteri ) with an average of 0.248 among 9 model species. Genetic differentiation in Korean native herbaceous plants estimated by ISSR marker in the present study (ФST=0.302) were lower than those of other foreign species (ФST=0.410).
In case of woody plants however, the level of genetic differentiation in Korean native plants was comparable or slightly higher than those of other species abroad (Ф ST=0.250). Based on allozyme marker, the genetic differentiation in Korean native plants estimated in this study (FST=0.248 for herbaceous plants) and previous studies (FST=0.239 for herbaceous plants, FST=0.076 for woody plants) is also similar with those of other species from abroad (FST=0.233 for herbaceous plants, FST=0.076 for woody plants).
Even within the same species, the extent of vegetative propagation was greatly different according to habitat environment. However, spatial extent by clonal propagation was considered to be greatly influenced by species-specific breeding system. In particular, species such as Wedelia prostrate that propagates by aboveground runner appeared to have larger spatial propagation range than species that propagates by rootstock. Of the species that propagate by rootstock, Epimedium koreanum or Scopolia japonica formed mostly small bunches without spreading spatially because of rhizome growing short.
For observation sites of most of the species for which spatial genetic structure was analyzed, spatial correlation was recognized. For woody plants, Sp values were between 0.006 (Betula ermanii ) and 0.102 (Abelia tyaihyoni ) with a mean of 0.027.
For perennial herbs, Sp values were between 0.002 (Lilium distichum) and 0.333 (Viola websteri ) with a mean of 0.088, three times higher than that of woody plants. In both woody plants and perennial herbs, vegetatively propagated species showed higher degree of spatial structure compared to sexually reproduced species.
2.3 Sampling strategy based on genetic diversity and ecological characteristics
The main objective of this study is to estimate the optimum sample size (i.e., number of populations and individuals) that secures the target amount of genetic variation in each species by comparing random sampling and preferential sampling based on genetic diversity and ecological characteristics of the model species. The optimum sample sizes were estimated by simulating the genetic data using the SGD program.
In the second place based on the results of the model species, we tried to propose an outline of sampling strategy for genetic conservation that can be applied to a number of forest plant species.
To provide the optimum sample sizes, three conservation goals for genetic diversity were applied. First, the sample size to capture on average either 90% or 95% of all alleles. Second, the sample size to capture all common alleles with 90% or 99.9% probabilities. Third, the sample size needed to obtain SI±10% (ISSR marker) and He±10% (allozyme marker) of the species level value with 95% or 99% probabilities.
The mean numbers of populations for achieving the three proposed conservation goals were between 4 and 10 by species, corresponding 40-100% of populations for the 14 species analyzed (6-14 populations). Thus, when species are to be preserved by randomly selecting populations, it is important to include most of the populations.
However, after the sampling that preferentially takes the populations with high genetic diversity, it was found that the conservation goal including 95% of all alleles on average was met by using just 60% of the populations for random sampling. Thus, for the efficient preservation of genetic resources, establishing a preservation strategy based on genetic diversity data, as much as possible, is desirable.
For finding a sampling method that can be applied to species for which genetic diversity data were unavailable, the genetic diversity parameters and correlation between the environmental and ecological characteristics were analyzed. This measure aims to identify factors that influence genetic diversity that showed correlation with latitude, longitude, soil, and vegetation characteristics of habitat by species, though factors common to multiple species were rare. However, distribution area and species richness were positively associated with genetic diversity parameters in many model species. Thus, the two characteristics were considered the most appropriate environmental parameter for predicting genetic diversity. Preferential sampling from nine species based on these two ecological characteristics showed that conservation goals were met with same or smaller number of populations (1 to 3) as was in random sampling.
The outline of sampling strategy resulted from the analysis of 22-model species for ensuring the conservation goals is as follows:
In the case of individual sampling, the optimum sample sizes were abstracted as follows.
- The sample size to capture on average 95% of all alleles was 230 individuals.
- The sample size to capture all common alleles with 99.9% probability was 130 individuals.
- The sample size to obtain SI±10% or He±10% of the value at the species level with 99% probability was 225 individuals.
Considering the aforementioned results, 250 individuals randomly chosen can cover all three conservation targets. This size can be applied as a useful standard to secure the representative amount of genetic diversity of a species. It would be reasonable and safe for ex-situ conservation measures such as field gene bank or for establishing a breeding population of a species.
In the case of population sampling, almost all of the population was needed to meet three conservation targets (especially for all alleles and all common alleles) due to genetic differentiation among populations. However, the sample size of 50 populations recommended by previous researchers is too large to be applied in Korea. Since most rare or endemic species in Korea remained in less than 15 populations, we recommended 15 as the practical number of populations for gene conservation.
In conclusion, 250 individuals should be sampled for the genetic conservation at the species level, even when sampling a number of populations is inapplicable. The sample size of 50 individuals per population is recommended when a number of populations are under consideration.
목차 Contents
- 표 지 ... 1
- 제출문 ... 2
- 보고서 요약서 ... 3
- 요약문 ... 4
- SUMMARY ... 20
- CONTENTS ... 26
- 목차 ... 27
- 제 1 장 연구개발과제의 개요 ... 28
- 1. 연구의 필요성 ... 28
- 2. 추진목표 ... 29
- 제 2 장 국내외 기술개발 현황 ... 31
- 1. 국외 연구동향 ... 31
- 2. 국내 연구동향 ... 32
- 제 3 장 연구개발수행 내용 및 결과 ... 34
- 제 1 절: 산림식물 보존전략 수립을 위한 분포특성 조사 ... 34
- 1. 서론 ... 34
- 2. 연구범위 및 방법 ... 35
- 3. 연구결과 ... 39
- 4. 고찰 ... 256
- 제 2 절. 산림식물 보존전략 수립을 위한 모델종의 유전다양성 및 구조 분석 ... 261
- 1. 서론 ... 261
- 2. 연구범위 및 방법 ... 262
- 3. 연구결과 ... 271
- 4. 고찰 ... 452
- 제 3 절: 유전다양성과 생태적 특성을 고려한 표본추출 전략 ... 466
- 1. 서론 ... 466
- 2. 연구배경 ... 468
- 3. 연구방법 ... 471
- 4. 연구결과 ... 474
- 5. 고찰 ... 504
- 6. 표본추출 기본전략 ... 506
- 제 4 장 목표달성도 및 관련분야에의 기여도 ... 513
- 1. 식물유형별 자생지환경특성분석 ... 513
- 2. 적정표본 추출을 위한 프로그램 개발 ... 513
- 3. 모델종의 유전변이 분석 ... 513
- 4. 생물적, 유전적 자료에 근거한 적정 보존 규모 결정 ... 513
- 5. 표본추출 지침 작성 ... 513
- 6. 연구성과 ... 514
- 제 5 장 연구개발결과의 활용계획 ... 518
- 1. 활용 계획 ... 518
- 2. 기대 효과 ... 518
- 3. 추가 연구의 필요성 ... 518
- 제 6 장 참고문헌 ... 519
- 제 7 장 첨부자료 ... 547
- 끝페이지 ... 746
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