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木麻黄(Casuarina spp.)具有速生、防风、固沙、抗逆及耐瘠薄等优良特性,是重要的防护林、用材林和多用途林树种[1-2]。我国引种木麻黄有110多年历史,最早于1897年引入台湾[3],20世纪80年代中期,借助国际合作项目,我国开始系统研究木麻黄种质资源的引种和选育[4]。目前,我国引进的木麻黄有20多种,人工种植面积达30多万公顷,但主要为短枝木麻黄(C. equisetifolia L.)、细枝木麻黄(C. cunninghamiana Miq.)和粗枝木麻黄(C. glauca Sieber. ex Spr.)。由于造林树种单一,种质资源匮乏,且长期受自然灾害及病虫害的影响,严重制约了木麻黄人工林的可持续经营[5]。开展山地木麻黄(C. junghuhniana Miq.)种质资源引种与测试,可丰富木麻黄种质资源,为木麻黄新品种创制和选育等提供基础材料,从而有利于提高沿海防护林的稳定性。
山地木麻黄原产印度尼西亚,具有固氮、速生、抗旱及耐水湿等特性,天然分布于山地及沿海地区,已被广泛引种到世界热带及亚热带地区。国外对山地木麻黄的研究主要集中在种源间生长表现和适应性比较以及优良种源选择[6-8],对其种源遗传变异规律及遗传参数估算研究较少[9]。我国引种山地木麻黄始于20世纪80年代,相继在沿海各地开展小规模引种试验。1996年以来,中国林科院热带林业研究所在广东湛江和福建漳州开展了国际山地木麻黄种源试验,对其遗传参数进行估算,并筛选出适合华南地区生长的优良种源[10-11];福建省林科院在福建漳州开展了滨海沙地山地木麻黄种源试验[12-13];但海南省尚无系统的山地木麻黄种源试验报道,特别是针对大量原产地种质材料的测试。本文以从澳大利亚引进的27个山地木麻黄种源为材料开展种源试验,分析其生长、形质等性状方面的变异状况,揭示种源间性状的遗传变异规律,同时应用坐标综合评定法对各种源间生长和形质性状进行综合评价,以期为山地木麻黄的良种选育和种质资源的合理利用提供参考。
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试验所用山地木麻黄27个种源种子均由澳大利亚林木种子中心提供(表 1)。2008年3月开始育苗,9月造林,采用机耕全垦方式整地,株行距2 m×2 m,造林时每株施150 g复合肥作为基肥。
表 1 27个山地木麻黄种源信息
Table 1. Details of seed origin of 27 Casuarina junghuhniana provenances
编号
No.种源号
Provenance code采种地点
Locality /Country纬度(S)
Latitude经度(E)
longitude海拔
Altitude/mJ1 18952 Mt Willis, East Java, Ind. 7°50’ 111°47’ 1 500 J2 18948 Mt Kawi, East Java, Ind. 7°55’ 112°25’ 2 000 J3 18951 Mt Arjuno, East Java, Ind. 7°45’ 112°35’ 1 350 J4 18950 Mt Bromo, East Java, Ind. 7°55’ 112°55’ 1 600 J5 18954 Mt Bromo, East Java, Ind. 7°55’ 112°55’ 2 500 J6 18949 Mt Agropuro, East Java, Ind 8°00’ 113°35’ 1 500 J7 18847 East Batu Kawu, Bali, Ind. 8°40’ 115°05’ 1 500 J8 18845 Mt Pohen, Bali, Ind. 8°40’ 115°05’ 2 000 J9 18844 Mt Tapak, Bali, Ind. 8°45’ 115°15’ 1 500 J10 18846 Mt Pengalongan, Bali, Ind. 8°50’ 115°15’ 1 500 J11 18849 Kintamani, Bali, Ind 8°13’ 115°20’ 1 500 J12 18848 Mt abang, Bali, Ind 8°55’ 115°25’ 1 500 J13 18850 Mt Santong, lombok, Ind. 8°25’ 116°28’ 1 500 J14 18851 Mt lamore, Lombok, Ind 8°25’ 116°45’ 1 500 J15 18852 Mt Tambora, Sumbawa, Ind 8°20’ 117°55’ 1 500 J16 19489 Kapan, Kupang, Timor, Ind. 10°13’ 123°38’ 600 J17 19490 Camplong, Timor, Ind 10°05’ 123°57’ 600 J18 17878 Noelmina river, Timor, Ind. 9°59’ 124°06’ 170 J19 17877 25 km SW Soe, Timor, Ind. 9°54’ 124°14’ 550 J20 19491 Buat, Soe, Timor, Ind. 9°51’ 126°16’ 800 J22 19239 Kari-Muguga, Ken. 1°16’ 36o36’ 2 060 J23 18953 Mt. Arjuno, East Java, Ind. 7°42’ 112o33’ 1 350 J24 19238 KEFRI Headquarters, Ken. 1°13’ 36°39’ 2 080 J25 19242 Kabiruini, Ken. 0°23’ 36°56’ 1 800 J26 19241 Thika, Ken. 1°02’ 37°12’ 1 440 J27 19237 Meru, Ken. 0°07’ 37°37’ 1 750 J28 18853 kwai Mission, Tanga, Tanz. 4°19’ 38°14’ 1 600 注:Ind.是印度尼西亚,Ken.是肯尼亚,Tanz.坦桑尼亚。
Note: Ind. is Indonesia, Ken. Is Kenyan. Tanz. is Tanzania.试验采用随机区组设计,以种源为处理,8次重复,每个种源采用单行6株小区。试验地四周用3行木麻黄无性系A8作为保护行。至2013年调查时,试验林未受任何人为和自然灾害的破坏,总体保存率达80%以上。由于2014年经历2次台风袭击(即威尔逊台风,14级,2014年7月18号登陆;海鸥台风,13级,2014年9月16号登陆),部分种源保存率较低。
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造林后2、5、7 a时,对所有种源进行每木调查,包括树高(H, m)、胸径(DBH, cm)和保存率(SUR, %)。抗风性观测在台风危害后进行。造林7 a时,对主干通直度(SFS)、主干分叉习性(AP)、侧枝直径(TPB)、侧枝密度(DPB)、侧枝长度(LPB)、侧枝分枝角(APB)、绿色小枝长度(LDB)、抗风性(RES)进行观测分级[14-16]
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山地木麻黄各性状以单株测定数据进行统计分析,单株材积(V, m3·株)计算公式为[17]:
V=3.141 592 6×DBH2×H/120 000
在方差分析前,对保存率进行反正弦转换,形质性状数据进行平方根转换。利用R语言结合软件ASReml 3.0和SPSS 22.0进行方差分析与多重比较、相关分析及遗传参数估算[18]。
观测值线性模型为:
$ {y_{ijk}} = \mu + {R_i} + {P_j} + R{P_{ij}} + {\eta _{ijk}} $
式中:yijk为观测值;μ为总体平均值;Ri为第i个区组的固定效应;Pj为第j个种源的随机效应;RPij为第i个区组与第j个种源互作的随机效应;ηijk为个体机误。
种源遗传力(Hp2)[19-20]:$H_p^2 = \frac{{\sigma _p^2}}{{\sigma _p^2 + \sigma _{pR}^2 + \sigma _p^2}}$
式中: σp2为种源方差分量,σpR2为种源与区组互作方差分量,σe2为误差方差分量。
表型方差分量(σph2)为:
$ \sigma _{ph}^2 = \sigma _p^2 + \sigma _{pR}^2 + \sigma _e^2 $
遗传变异系数(GCVA, %)为:
$ GC{V_A} = \frac{{{\sigma _p}}}{{\bar X}} \times 100{\rm{\% }} $
式中:X是性状的平均值,σp是性状种源方差分量的平方根。
性状间的遗传相关系数(rG)和表型相关系数(rP)为:
$ {r_{G(xy)}} = \frac{{{{{\mathop{\rm cov}} }_c}(x, y)}}{{\sqrt {\sigma _{{\rm{cx}}}^2 \cdot \sigma _{cy}^2} }} $
$ {r_{P(xy)}} = \frac{{{{{\mathop{\rm cov}} }_p}(x, y)}}{{\sqrt {\sigma _{px}^2 \cdot \sigma _{py}^2} }} $
式中:CovG(x, y)为性状x与性状y间的遗传协方差,σGx2为性状x的遗传方差分量,σGy2为性状y的遗传方差分量;Covp(x, y)为性状x与性状y间的表型协方差,σpx2为性状x的表型方差分量,σpy2为性状y的表型方差分量。
种源选择遗传增益估算:
$ {\rm{\Delta }}G = H_p^2 \cdot S \cdot {\bar X^{ - 1}} \times 100{\rm{\% }} $
式中:ΔG为种源性状遗传增益;Hp2为种源性状遗传力;S为入选种源性状均值与种源性状总均值的离差;X-1为种源性状总均值。
基于各性状的观测值,采用坐标综合评定法[21]对各种源进行综合评价,筛选优良种源。
$ {P_i} = \sqrt {\sum\limits_{j = 1}^n {{{\left( {1 - \frac{{{X_{ij}}}}{{{X_{j\max }}}}} \right)}^2}} } $
式中:Pi为综合值, Xij为第i个种源第j个性状的平均观测值, Xjmax为第j个性状的最大平均观测值,n为性状个数。
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表 2表明:在测定的3个年份,山地木麻黄各种源间保存率均差异显著(P < 0.05);造林后2、5、7 a,其保存率分别为19.67%~99.41%、0.59%~93.93%、0.28%~76.67%,其中,造林后2、5 a时,山地木麻黄的整体平均保存率在74%以上,7 a时因遭受超强台风袭击,整体保存率降至40%。造林后2、5 a时,种源18948、19489和17878保存率最高,均达90%以上;造林后7 a时,种源18844、18846、18849和19489保存率最高,均在70%以上。
表 2 27个山地木麻黄种源的保存率和抗风性
Table 2. Average surivival values and wind-resistance score with standard error of 27 Casuarina junghuhniana provenances
种源
Provenance保存率Survival /% 抗风性
Wind-resistance2 a 5 a 7 a 18952 88.89±0.61ab 65.25±0.39ab 9.22±1.00efg 1.31±0.15e 18948 96.72±0.83a 93.44±1.02a 38.85±2.63abcdef 2.07±0.21cd 18951 98.33±0.75a 89.56±1.58a 13.65±1.46defg 1.55±0.19de 18950 95.80±1.06ab 85.87±0.92a 37.19±2.70abcdef 3.13±0.17a 18954 94.59±0.84ab 80.11±0.66ab 61.83±1.52ab 3.40±0.17a 18949 97.53±0.59a 58.76±4.56ab 13.65±1.93defg 1.83±0.23de 18847 19.67±5.29c 0.59±0.59c 0.28±0.28g 2.50±1.50bc 18845 93.44±1.02ab 73.26±3.86ab 52.44±3.03abcd 3.46±0.15a 18844 98.90±0.47a 86.35±3.62a 73.26±3.86a 3.24±0.17a 18846 99.41±0.59a 88.89±0.61a 76.67±1.27a 3.40±0.12a 18849 96.72±0.83a 89.33±1.14a 72.64±1.12a 3.60±0.17a 18848 94.59±0.84ab 68.49±3.33ab 24.88±5.79bcdef 2.10±0.21cd 18850 87.78±1.26ab 65.25±2.47ab 51.43±1.72abcd 3.13±0.16a 18851 93.25±0.59ab 37.19±4.39b 6.07±1.62fg 1.67±0.27de 18852 88.89±0.61ab 70.10±0.73ab 59.07±0.46abc 3.31±0.18a 19489 97.53±0.59a 91.98±0.76a 74.79±1.24a 3.27±0.19a 19490 98.33±0.75a 74.52±3.10ab 56.66±2.88abc 3.25±0.17a 17878 98.33±0.75a 93.93±1.73a 69.52±0.40a 3.00±0.19ab 17877 91.98±0.76ab 89.33±1.14a 63.51±2.73ab 3.28±0.20a 19491 72.64±5.33b 62.84±4.19ab 21.05±1.83bcdefg 1.56±0.18de 19239 85.87±0.92ab 70.45±3.16ab 56.66±2.98abc 3.30±0.12a 18953 94.91±1.35ab 84.86±1.75a 16.93±1.82cdefg 1.77±0.20de 19238 83.62±0.45ab 83.62±0.45a 46.56±1.70abcde 3.28±0.18a 19242 93.25±0.59ab 67.55±2.54ab 35.52±1.47abcdef 3.14±0.17a 19241 85.65±0.82ab 61.83±1.15ab 41.54±0.21abcdef 3.00±0.11a 19237 97.53±0.59a 80.66±1.22ab 38.17±1.05abcdef 2.81±0.18ab 18853 82.02±0.62ab 60.85±0.22ab 41.24±0.44abcdef 3.40±0.23a 平均Means 92.07±0.05 74.09±0.09 40.90±0.09 2.74±0.04 F test ** ** ** ** 注:* P<0.05的显著水平,** P<0.01的极显著水平,ns为不显著,小写字母表示在0.05水平上显著,下同。
Note: * P<0.05 level of significance, ** P<0.01 level of significance, ns-no significant, lowercase letter indicate significant at 0.05 level, The same below.表 2还表明:造林后7 a时,山地木麻黄种源抗风性差异显著,种源18952的抗风性最差,种源18849、18845、18954、18846、18852、19239和18853的抗风性相对较好。
由于超强台风的影响,部分种源保存率较低,树干倒伏、折断较严重,小区数据缺失,为保证分析结果的可靠性,基于上述保存率及抗风性分析,剔除7 a时缺失5个小区以上的9个种源(18952、18948、18951、18949、18847、18848、18851、19491和18953),仅对保存有5个小区以上的18个种源的生长和形质性状进行统计分析。
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从表 3可看出:造林后2、5、7 a时,18个种源的树高、胸径、单株材积均存在极显著差异(P < 0.01),说明生长性状在种源水平上具有很大的遗传改良潜力;各种源的树高均值分别为3.48~6.12、4.98~8.26、7.50~11.67 m,胸径均值分别为2.66~4.75、4.84~7.73、5.96~9.55 cm,单株材积均值分别为1.02×10-3~4.46×10-3、4.43×10-3~16.14×10-3、10.20×10-3~34.88×10-3 m3,其中,种源19490和17877表现最好,种源18846和18845表现最差。
表 3 造林后2、5、7 a时山地木麻黄18个种源树高、胸径及单株材积的均值及其多重比较结果
Table 3. Average values on height, DBH and individual volume with standard error and Duncan Multiple Range Test of 18 C. junghuhniana provenances at 2, 5 and 7 years after planting
种源
Provenance树高Height /m 胸径DBH/cm 单株材积Volume/×10-3m3 2 a 5 a 7 a 2 a 5 a 7 a 2 a 5 a 7 a 18950 4.54±0.21cdef 6.38±0.29cd 9.25±0.64cdef 3.17±0.21g 5.39±0.36cde 6.89±0.58efg 1.51±0.34def 6.14±1.29def 14.35±3.9def 18954 4.45±0.21def 6.55±0.30cd 9.95±0.54bcde 3.32±0.21defg 5.41±0.36cde 7.37±0.50efg 1.66±0.34cdef 6.51±1.33def 17.38±3.30def 18845 4.90±0.22bcde 5.94±0.32def 8.34±0.57efg 3.76±0.21cdef 5.17±0.39de 6.36±0.53fg 2.59±0.34b 4.96±1.42f 10.44±3.49f 18844 5.05±0.21bc 6.64±0.30cd 8.51±0.51defg 4.09±0.20bc 6.00±0.36bcd 6.59±0.47efg 2.95±0.33b 7.63±1.33cdef 11.14±3.08ef 18846 4.58±0.21cde 6.18±0.29de 8.15±0.49fg 3.34±0.20defg 4.84±0.35e 5.96±0.45g 1.75±0.33cdef 4.98±1.26f 10.20±2.99f 18849 5.03±0.21bcd 7.68±0.30ab 10.82±0.50abc 3.22±0.21g 6.35±0.36bc 7.84±0.46def 1.65±0.33cdef 9.45±1.33cde 19.01±3.04cdef 18850 5.32±0.22b 7.21±0.35bc 10.06±0.57bcd 3.84±0.22cd 6.73±0.42ab 8.14±0.52abcde 2.70±0.35b 9.75±1.52cde 18.47±3.49def 18852 4.64±0.22cde 6.35±0.32cd 8.81±0.55defg 3.27±0.22efg 5.45±0.39cde 6.96±0.50efg 1.73±0.35cdef 5.99±1.42ef 13.64±3.36def 19489 5.45±0.21b 7.92±0.28ab 11.46±0.49ab 4.20±0.21bc 7.59±0.34a 9.18±0.45abcd 2.91±0.33b 14.26±1.25ab 28.33±2.99abc 19490 6.12±0.21a 7.91±0.32ab 11.67±0.55a 4.75±0.20a 7.66±0.38a 9.55±0.51a 4.46±0.33a 15.86±1.38a 34.00±3.36a 17878 4.36±0.21ef 6.61±0.29cd 9.66±0.51cdef 3.36±0.20defg 6.42±0.35bc 8.01±0.46bcde 1.77±0.33cdef 10.03±1.26cde 22.97±3.13bcd 17877 5.96±0.22a 8.26±0.29a 11.45±0.53ab 4.44±0.21ab 7.73±0.36a 9.40±0.48abc 3.87±0.34a 16.14±1.29a 34.88±3.24a 19239 4.46±0.23def 6.58±0.34cd 9.00±0.55defg 3.79±0.23cde 6.46±0.41bc 7.65±0.51ef 2.42±0.36bc 10.10±1.49cd 20.87±3.36cde 19238 4.50±0.23cdef 6.74±0.29cd 10.17±0.58abcd 3.24±0.22fg 6.40±0.36bc 8.96±0.54abcd 1.85±0.36cde 11.43±1.29bc 31.45±3.56ab 19242 4.36±0.21ef 4.98±0.33g 7.61±0.66g 3.40±0.21defg 4.89±0.40de 6.72±0.61efg 2.30±0.34bcd 4.43±1.44f 12.67±4.00def 19241 3.48±0.23g 5.16±0.35fg 8.13±0.64fg 2.66±0.22h 5.13±0.42de 7.18±0.59efg 1.02±0.36f 5.94±1.54ef 19.35±3.90cdef 19237 3.95±0.21fg 5.47±0.31efg 7.50±0.66g 2.96±0.21gh 5.16±0.37de 7.25±0.59efg 1.23±0.33ef 4.87±1.36f 13.51±4.00def 18853 4.40±0.23ef 6.50±0.35cd 9.38±0.64cdef 3.19±0.23g 5.64±0.42bcde 7.36±0.59efg 1.85±0.36cde 7.71±1.52cdef 18.12±3.90def 均值Means 4.77±0.06 6.66±0.08 9.55±0.14 3.57±1.46 6.06±0.09 7.67±0.13 2.25±0.09 8.82±0.35 19.76±0.87 F test ** ** ** ** ** ** ** ** ** 表 4表明:造林后7 a时,山地木麻黄侧枝直径(TPB)、侧枝分枝角(APB)、绿色小枝长度(LDB)及主干通直度(SFS)在种源间差异极显著;主干分叉习性(AP)在种源间差异显著;而抗风性(RES)、侧枝密度(DPB)及侧枝长度(LPB)在种源间差异不显著。
表 4 造林后7 a时18个种源山地木麻黄形质性状的得分均值及其多重比较结果
Table 4. The mean score of qualitative traits with standard error and Duncan Multiple Range Test of 18 C. junghuhniana provenances at 7years after planting
种源
ProvenanceRES DPB TPB APB LPB LDB AP SFS 18950 3.13±0.06abc 3.33±0.05abc 3.25±0.08a 1.42±0.06bcd 1.92±0.03a 1.33±0.06bcd 4.79±0.08abcd 2.82±0.03efg 18954 3.40±0.05abc 3.06±0.05bc 3.00±0.07abcd 1.38±0.05bcd 1.94±0.03a 1.56±0.05abc 4.61±0.08abcd 3.29±0.03ab 18845 3.46±0.06ab 3.22±0.06bc 3.11±0.09ab 1.22±0.07d 2.00±0.04a 1.67±0.07ab 4.00±0.09cd 2.84±0.04efg 18844 3.24±0.05abc 3.45±0.04ab 3.05±0.06abcd 1.32±0.04cd 1.95±0.03a 1.64±0.04ab 4.58±0.08abcd 2.89±0.04def 18846 3.40±0.05ab 3.44±0.05ab 3.06±0.07abc 1.19±0.05d 2.00±0.03a 1.50±0.05abc 4.16±0.08bcd 3.20±0.03bcd 18849 3.60±0.05a 3.20±0.04bc 2.70±0.06abcde 1.30±0.05cd 2.00±0.03a 1.35±0.05bcd 4.21±0.08bcd 3.56±0.03a 18850 3.13±0.05abc 3.06±0.05bc 2.44±0.07bcde 1.44±0.05abcd 1.94±0.03a 1.44±0.05abc 3.85±0.09d 3.09±0.04bcde 18852 3.31±0.05abc 3.26±0.04bc 2.95±0.06abcd 1.47±0.05abcd 1.89±0.03a 1.32±0.05bcd 4.39±0.08abcd 3.08±0.04bcde 19489 3.27±0.05abc 3.43±0.04ab 2.62±0.06abcde 1.71±0.05ab 1.95±0.03a 1.67±0.04ab 4.91±0.08abc 3.20±0.03bcd 19490 3.25±0.05abc 3.31±0.05abc 2.38±0.07de 1.50±0.05abcd 2.00±0.03a 1.69±0.05ab 4.49±0.08abcd 3.03±0.04bcde 17878 3.00±0.05bc 3.29±0.04abc 2.43±0.06cde 1.81±0.05a 1.86±0.03a 1.00±0.04d 4.23±0.08abcd 2.63±0.03fg 17877 3.28±0.05abc 3.50±0.04ab 2.70±0.06abcde 1.75±0.05ab 1.95±0.03a 1.80±0.05a 4.95±0.08ab 3.15±0.03bcde 19239 3.30±0.06abc 3.26±0.04bc 2.87±0.06abcde 1.30±0.04cd 2.00±0.03a 1.61±0.04ab 4.75±0.08abcd 3.06±0.04bcde 19238 3.28±0.06abc 3.16±0.04bc 2.32±0.06e 1.47±0.05abcd 1.95±0.03a 1.37±0.05bcd 4.97±0.08ab 3.05±0.03bcde 19242 3.14±0.06abc 3.33±0.05abc 2.67±0.08abcde 1.67±0.06abc 2.00±0.03a 1.00±0.06d 4.34±0.08abcd 2.51±0.04g 19241 3.00±0.06abc 3.44±0.05ab 2.94±0.07abcde 1.38±0.05bcd 2.00±0.03a 1.31±0.05bcd 4.35±0.08abcd 2.89±0.04cdef 19237 2.81±0.06c 3.79±0.05a 3.14±0.07ab 1.14±0.06d 1.86±0.03a 1.43±0.05abc 5.11±0.08a 2.86±0.04def 18853 3.40±0.06abc 3.31±0.05c 2.69±0.08bcde 1.46±0.06d 1.92±0.03a 1.38±0.06cd 5.00±0.09ab 3.26±0.04abc 均值Means 3.25±0.01 3.32±0.01 2.77±0.02 1.45±0.01 1.95±0.01 1.46±0.01 4.54±0.02 3.03±0.01 F test ns ns ** ** ns ** * ** -
由表 5可知:树高、胸径、单株材积等生长性状的种源遗传力随着林龄的增长,呈先增长后降低的趋势,其种源遗传力分别为13.80%~16.00%、8.64%~11.90%、8.42%~14.30%。造林后7 a时,各形质性状间种源遗传力差异较大,TPB、APB、LDB、SFS的种源遗传力均>5%,RES、DPB、LPB、AP的种源遗传力均<1%。树高的遗传变异系数随着林龄的增加变化不大,其遗传变异系数为12%左右;胸径和单株材积的遗传变异系数随林龄的增加呈先增加后降低的趋势,其遗传变异系数分别为11.67%~13.67%、30.20%~38.11%。造林后7 a时,除DPB的遗传变异系数接近于0,其他形质性状的遗传变异系数为0.34%~5.56%。采用20%的入选率,造林后2、5、7 a时,树高、胸径、单株材积的遗传增益分别为2.59%~3.08%、1.95%~2.69%、4.86%~9.08%。造林后7 a时,形质性状的遗传增益为3.29×10-8%~2.45%,明显低于生长性状,选择生长性状进行遗传改良效果较好。
表 5 山地木麻黄各性状的方差分量(种源、互作、表型和误差)、种源遗传力、遗传变异系数及遗传增益
Table 5. Estimates of variances (σp2, σp2R, σph2, σe2) for additive, interaction, phenotypic and error, respectively, and Provenance heritability (HP2), genetic coefficient of variation (GCVA) and genetic gains (ΔG) for various traits
性状Trait σp2 σp2R σph2 σp2e Hp2/% GCVA/% ΔG/% H2 0.344 0.581 2.339 1.414 14.70** 12.30 2.92 DBH2 0.182 0.767 2.104 1.155 8.64* 11.96 1.95 V2 0.461a 2.590a 5.481a 2.430a 8.42* 30.20 4.86 H5 0.666 0.648 4.164 2.850 16.00** 12.25 3.08 DBH5 0.685 0.846 5.778 4.247 11.90** 13.67 2.69 V5 0.011b 0.012b 0.079b 0.055b 14.30** 38.11 9.08 H7 1.290 1.550 9.370 6.530 13.80** 11.89 2.59 DBH7 0.802 1.199 7.734 5.733 10.40* 11.67 2.17 V7 0.047b 0.061b 0.355b 0.247b 13.40** 34.85 8.41 RES 0.439b 0.002 0.080 0.077 0.50ns 0.64 0.03 DPB 0.016a 0.003 0.033 0.030 0.05a** 3.84a 3.29b TPB 0.005 0.003 0.077 0.069 6.88* 2.62 0.91 APB 0.004 0.003 0.047 0.040 8.75* 4.41 1.70 LPB 0.043b 0.002 0.014 0.012 0.30 ns 0.34 0.01 LDB 0.007 0.001 0.046 0.039 14.30** 5.56 2.45 AP 0.002 0.019 0.248 0.227 0.72 ns 0.93 0.07 SFS 0.003 0.009 0.051 0.038 6.54* 1.91 0.65 注:表中a表示方差分量×10-3, b表示方差分量×10-6;H2、DBH2、V2、H5、DBH5、V5、H7、DBH7、V7分别表示造林后2、5、7 a时的树高、胸径和单株材积;RES、DPB、TPB、APB、LPB、LDB、AP、SFS为造林后7 a时测定;下同
Note: a represents variances×10-3, b represents variances×10-6. H2, DBH2, V2, H5, DBH5, V5, H7, DBH7, V7 represents the height, diameter at breast height and volume at 2, 5 and 7 years after planting, respectively. RES, DPB, TPB, APB, LPB, LDB, AP, SFS is measured at 7 years after planting. The same below.由表 6可看出:树高、胸径、单株材积3个生长性状间的表型和遗传均呈极显著相关,相关系数均>0.68。TPB与3个生长性状间的遗传和表型均呈显著或极显著负相关,而APB与3个生长性状间的遗传和表型均呈显著或极显著正相关;APB除与TPB间的表型和遗传呈极显著负相关外,与其他形质性状均相关不显著;树高与SFS间的表型和遗传呈极显著正相关,这对选择生长快和干形好的种源极为有利。不同林龄间,树高和胸径的遗传相关系数均在0.78以上,达极显著相关,说明对生长性状进行早期选择是可行的。
表 6 山地木麻黄各性状的遗传相关系数(上三角),表型相关系数(下三角)
Table 6. Genotypic (upper triangle), phenotypic correlations (lower triangle) for various traits
H2 H5 H7 DBH2 DBH5 DBH7 V2 V5 V7 TPB APB LDB AP SFS H2 0.97** 0.94** 0.94** 0.87** 0.82** 0.92** 0.86** 0.75** -0.52* 0.45* 0.77** -0.17ns 0.38* H5 0.83** 0.99** 0.84** 0.91** 0.85** 0.84** 0.89** 0.78** -0.63** 0.51* 0.69** 0.10ns 0.66** H7 0.77** 0.87** 0.82** 0.78** 0.89** 0.81** 0.74** 0.94** -0.76** 0.67** 0.50* 0.26ns 0.60** DBH2 0.88** 0.74** 0.68** 0.88** 0.84** 0.87** 0.93** 0.80** -0.52* 0.53* 0.86** -0.02ns 0.11ns DBH5 0.77** 0.86** 0.94** 0.81** 0.93** 0.90** 0.94** 0.85** -0.90** 0.76** 0.62** 0.37ns 0.35ns DBH7 0.75** 0.83** 0.82** 0.77** 0.99** 0.85** 0.89** 0.98** -0.92** 0.81** 0.42* 0.75* 0.28ns V2 0.86** 0.76** 0.68** 0.99** 0.82** 0.77** 0.91** 0.84** -0.54* 0.51* 0.80** 0.01ns 0.06ns V5 0.72** 0.85** 0.74** 0.74** 0.99** 0.99** 0.82** 0.89** -0.84** 0.69** 0.61** 0.47ns 0.36ns V7 0.70** 0.79** 0.72** 0.72** 0.96** 0.94** 0.77** 0.99** -0.88** 0.75** 0.40ns 0.75* 0.25ns TPB -0.43** -0.50** -0.55** -0.52* -0.56** -0.60** -0.47** -0.48** -0.55** -0.82** 0.15ns -0.06ns -0.07ns APB 0.23** 0.28** 0.30** 0.23** 0.30** 0.29** 0.20** 0.24** 0.25** -0.15* -0.25ns 0.11ns -0.42ns LDB 0.15* 0.11* 0.09ns 0.16* 0.03ns -0.02ns 0.15* 0.07ns 0.04ns 0.27** - 0.35ns 0.55* AP 0.04ns 0.04ns 0.05ns 0.06* -0.04ns -0.03ns 0.06ns - 0.03ns -0.07ns 0.01ns 0.07ns -0.16ns SFS 0.15** 0.20** 0.17** 0.05ns 0.07* 0.05ns 0.07ns 0.11* 0.10* -0.10* -0.09ns 0.07ns 0.25** -
选择种源间差异显著的生长及形质性状,应用坐标综合评定法对18个种源进行综合评定(表 7)发现,最好的3个种源为17877、19489和19490,其评估得分分别为0.074 5,0.095 4和0.212 2;种源19242最差,其评估得分值为1.277 0。
表 7 参试18个种源综合排名
Table 7. Ordinal ranking of 18 C. junghuhniana provenances
种源
ProvenanceH7 D7 V7 SUR TPB APB LDB AP SFS 综合评价
Overview排名
Rank18950 0.042 9 0.077 9 0.346 5 0.265 2 0.000 0 0.047 1 0.067 2 0.000 0 0.043 5 0.890 3 14 18954 0.021 7 0.052 1 0.251 7 0.025 3 0.005 9 0.057 7 0.017 4 0.001 5 0.005 8 0.439 1 5 18845 0.081 3 0.111 9 0.491 1 0.039 4 0.001 8 0.105 3 0.005 5 0.027 5 0.040 8 0.904 7 15 18844 0.073 2 0.095 9 0.463 2 0.002 0 0.004 0 0.073 7 0.008 3 0.011 0 0.035 3 0.766 5 13 18846 0.091 0 0.141 7 0.500 5 0.000 0 0.003 3 0.118 2 0.027 8 0.034 9 0.010 2 0.927 5 16 18849 0.005 3 0.032 1 0.207 1 0.002 8 0.028 6 0.079 3 0.062 5 0.031 6 0.000 0 0.449 3 6 18850 0.019 0 0.021 9 0.221 4 0.108 4 0.062 5 0.042 3 0.040 6 0.061 3 0.017 6 0.594 8 9 18852 0.059 8 0.073 4 0.370 9 0.052 7 0.008 7 0.034 4 0.072 4 0.020 1 0.018 2 0.710 6 10 19489 0.000 3 0.001 5 0.035 2 0.000 6 0.037 7 0.002 8 0.005 5 0.001 6 0.010 2 0.095 4 2 19490 0.000 0 0.000 0 0.000 6 0.068 1 0.072 5 0.029 3 0.003 9 0.015 1 0.022 7 0.212 2 3 17878 0.029 5 0.026 2 0.116 5 0.008 7 0.063 9 0.000 0 0.197 5 0.030 2 0.069 4 0.542 0 8 17877 0.000 4 0.000 2 0.000 0 0.029 5 0.028 6 0.001 1 0.000 0 0.001 0 0.013 7 0.074 5 1 19239 0.052 2 0.039 7 0.161 3 0.085 6 0.013 7 0.077 9 0.011 3 0.005 1 0.020 4 0.467 3 7 19238 0.016 5 0.003 9 0.009 7 0.154 2 0.082 6 0.034 4 0.057 5 0.000 8 0.020 9 0.380 5 4 19242 0.121 2 0.087 8 0.405 4 0.317 0 0.032 2 0.006 2 0.197 5 0.022 8 0.086 8 1.277 0 18 19241 0.092 2 0.061 7 0.198 1 0.209 9 0.009 2 0.057 7 0.073 4 0.022 3 0.035 6 0.760 0 11 19237 0.127 6 0.058 1 0.375 3 0.252 2 0.001 1 0.135 7 0.042 6 0.000 0 0.039 3 1.031 8 17 18853 0.038 6 0.052 7 0.230 8 0.312 6 0.029 4 0.037 0 0.053 3 0.000 5 0.007 4 0.762 3 12
山地木麻黄种源在海南临高的遗传变异及选择
Genetic Variation and Selection of Casuarina junghuhniana Provenances at Lingao, Hainan
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摘要:
目的 研究山地木麻黄种源间抗风性、生长及形质性状的遗传变异规律,为山地木麻黄的良种选育和种质资源的合理开发利用提供科学依据。 方法 以27个山地木麻黄种源为试验材料,于造林后2、5、7 a时测定山地木麻黄种源的树高、胸径、单株材积和保存率等数量性状,并于造林后7 a时调查主干分叉习性(AP)、主干通直度(SFS)、侧枝密度(DPB)、侧枝直径(TPB)、绿色小枝长度(LDB)、侧枝分枝角(APB)、侧枝长度(LPB)等形质性状以及抗风性(RES),通过方差分析、相关性分析及遗传参数估算揭示其遗传变异规律。应用坐标综合评定法对山地木麻黄种源进行综合评定。 结果 表明:造林后2、5、7 a时,27个山地木麻黄种源间保存率和抗风性差异显著(P < 0.05);对造林后7 a时保存率较高的18个种源进一步分析显示,上述3个年份各种源间树高、胸径和单株材积等生长性状均存在极显著差异(P < 0.01);7 a时,TPB、APB、LDB、AP和SFS等形质性状在种源间亦存在显著或极显著差异;生长性状的种源遗传力明显高于形质性状,二者分别受中度或中度偏下和低度遗传控制;随着林龄的增长,树高的遗传变异系数变化不大,而胸径和单株材积的遗传变异系数呈先增加后降低的趋势,树高、胸径、单株材积的遗传变异系数分别为11.89%~12.30%、11.67%~13.67%、30.20%~38.11%;7 a时,形质性状的遗传变异系数为3.84×10-5%~5.56%。性状间相关分析表明:树高作为山地木麻黄早期选择性状较适宜。 结论 依据坐标综合评定法,筛选出17877、19489和19490等3个优良种源,可在生产上大面积推广。 Abstract:Objective In order to screen out superior provenances and individual trees for further cross-breeding, the inter-provenance genetic variations in terms of wind-resistance, growth and morphological traits were discussed in this context due to a trial of Casuarina junghuhniana including 27 provenances at Lingao, Hainan, China. Method The height, DBH, volume and survival were measured at the 2nd, 5th and 7th years after planting, and the qualitative traits including AP, SFS, DPB, TPB, LDB, APB, LPB and RES were investigated at the 7th year after planting. The genetic variations of these traits were studied by variance analysis, genotypic and phenotypic correlations, and genetic parameters. The provenances of C. junghuhniana were also assessed by comprehensive coordinate method. Result Significant differences (P < 0.05) were detected among 27 provenances in survival and RES at the 2nd, 5th and 7th years after planting. For a further analysis, the top 18 provenances in higher survivals at age of 7 indicated signi? cant differences (P < 0.01) in tree height, DBH, volumes among provenances at the 2nd, 5th and 7th years after planting. It was also found that there were significant differences in TPB, APB, LDB, SFS (P < 0.01) and AP (P < 0.05) among provenances 7 years after planting. The heritability was moderate for growth traits and low for qualitative traits, suggesting the genetic controls were moderate on growth and weak on qualitative traits. Trends of coefficient of genetic variation for height were nearly stable, while the coefficients of genetic variations in DBH and volume tended to decreasing at initial and then increasing with age. The coefficients of genetic variation of height, DBH and volume were 11.89%-12.30%, 11.67%-13.67% and 30.20%-38.11%, respectively, and those for qualitative traits 7 years after planting ranged from 3.84×10-5% to 5.56%. The high age-age and trait-trait phenotypic and genetic correlations for height and DBH indicated that early selection and multiple traits selection were feasible. Height was superior to other traits in the early selection for C. junghuhniana due to its genetic stability. Conclusion By analyzing nine traits at the 7th year after planting by comprehensive coordinate method, three optimal provenances were selected, which are worthy to be developed for producing and cross-breeding. -
Key words:
- Casuarina junghuhniana
- / provenance selection
- / heritability
- / genetic variations
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表 1 27个山地木麻黄种源信息
Table 1. Details of seed origin of 27 Casuarina junghuhniana provenances
编号
No.种源号
Provenance code采种地点
Locality /Country纬度(S)
Latitude经度(E)
longitude海拔
Altitude/mJ1 18952 Mt Willis, East Java, Ind. 7°50’ 111°47’ 1 500 J2 18948 Mt Kawi, East Java, Ind. 7°55’ 112°25’ 2 000 J3 18951 Mt Arjuno, East Java, Ind. 7°45’ 112°35’ 1 350 J4 18950 Mt Bromo, East Java, Ind. 7°55’ 112°55’ 1 600 J5 18954 Mt Bromo, East Java, Ind. 7°55’ 112°55’ 2 500 J6 18949 Mt Agropuro, East Java, Ind 8°00’ 113°35’ 1 500 J7 18847 East Batu Kawu, Bali, Ind. 8°40’ 115°05’ 1 500 J8 18845 Mt Pohen, Bali, Ind. 8°40’ 115°05’ 2 000 J9 18844 Mt Tapak, Bali, Ind. 8°45’ 115°15’ 1 500 J10 18846 Mt Pengalongan, Bali, Ind. 8°50’ 115°15’ 1 500 J11 18849 Kintamani, Bali, Ind 8°13’ 115°20’ 1 500 J12 18848 Mt abang, Bali, Ind 8°55’ 115°25’ 1 500 J13 18850 Mt Santong, lombok, Ind. 8°25’ 116°28’ 1 500 J14 18851 Mt lamore, Lombok, Ind 8°25’ 116°45’ 1 500 J15 18852 Mt Tambora, Sumbawa, Ind 8°20’ 117°55’ 1 500 J16 19489 Kapan, Kupang, Timor, Ind. 10°13’ 123°38’ 600 J17 19490 Camplong, Timor, Ind 10°05’ 123°57’ 600 J18 17878 Noelmina river, Timor, Ind. 9°59’ 124°06’ 170 J19 17877 25 km SW Soe, Timor, Ind. 9°54’ 124°14’ 550 J20 19491 Buat, Soe, Timor, Ind. 9°51’ 126°16’ 800 J22 19239 Kari-Muguga, Ken. 1°16’ 36o36’ 2 060 J23 18953 Mt. Arjuno, East Java, Ind. 7°42’ 112o33’ 1 350 J24 19238 KEFRI Headquarters, Ken. 1°13’ 36°39’ 2 080 J25 19242 Kabiruini, Ken. 0°23’ 36°56’ 1 800 J26 19241 Thika, Ken. 1°02’ 37°12’ 1 440 J27 19237 Meru, Ken. 0°07’ 37°37’ 1 750 J28 18853 kwai Mission, Tanga, Tanz. 4°19’ 38°14’ 1 600 注:Ind.是印度尼西亚,Ken.是肯尼亚,Tanz.坦桑尼亚。
Note: Ind. is Indonesia, Ken. Is Kenyan. Tanz. is Tanzania.表 2 27个山地木麻黄种源的保存率和抗风性
Table 2. Average surivival values and wind-resistance score with standard error of 27 Casuarina junghuhniana provenances
种源
Provenance保存率Survival /% 抗风性
Wind-resistance2 a 5 a 7 a 18952 88.89±0.61ab 65.25±0.39ab 9.22±1.00efg 1.31±0.15e 18948 96.72±0.83a 93.44±1.02a 38.85±2.63abcdef 2.07±0.21cd 18951 98.33±0.75a 89.56±1.58a 13.65±1.46defg 1.55±0.19de 18950 95.80±1.06ab 85.87±0.92a 37.19±2.70abcdef 3.13±0.17a 18954 94.59±0.84ab 80.11±0.66ab 61.83±1.52ab 3.40±0.17a 18949 97.53±0.59a 58.76±4.56ab 13.65±1.93defg 1.83±0.23de 18847 19.67±5.29c 0.59±0.59c 0.28±0.28g 2.50±1.50bc 18845 93.44±1.02ab 73.26±3.86ab 52.44±3.03abcd 3.46±0.15a 18844 98.90±0.47a 86.35±3.62a 73.26±3.86a 3.24±0.17a 18846 99.41±0.59a 88.89±0.61a 76.67±1.27a 3.40±0.12a 18849 96.72±0.83a 89.33±1.14a 72.64±1.12a 3.60±0.17a 18848 94.59±0.84ab 68.49±3.33ab 24.88±5.79bcdef 2.10±0.21cd 18850 87.78±1.26ab 65.25±2.47ab 51.43±1.72abcd 3.13±0.16a 18851 93.25±0.59ab 37.19±4.39b 6.07±1.62fg 1.67±0.27de 18852 88.89±0.61ab 70.10±0.73ab 59.07±0.46abc 3.31±0.18a 19489 97.53±0.59a 91.98±0.76a 74.79±1.24a 3.27±0.19a 19490 98.33±0.75a 74.52±3.10ab 56.66±2.88abc 3.25±0.17a 17878 98.33±0.75a 93.93±1.73a 69.52±0.40a 3.00±0.19ab 17877 91.98±0.76ab 89.33±1.14a 63.51±2.73ab 3.28±0.20a 19491 72.64±5.33b 62.84±4.19ab 21.05±1.83bcdefg 1.56±0.18de 19239 85.87±0.92ab 70.45±3.16ab 56.66±2.98abc 3.30±0.12a 18953 94.91±1.35ab 84.86±1.75a 16.93±1.82cdefg 1.77±0.20de 19238 83.62±0.45ab 83.62±0.45a 46.56±1.70abcde 3.28±0.18a 19242 93.25±0.59ab 67.55±2.54ab 35.52±1.47abcdef 3.14±0.17a 19241 85.65±0.82ab 61.83±1.15ab 41.54±0.21abcdef 3.00±0.11a 19237 97.53±0.59a 80.66±1.22ab 38.17±1.05abcdef 2.81±0.18ab 18853 82.02±0.62ab 60.85±0.22ab 41.24±0.44abcdef 3.40±0.23a 平均Means 92.07±0.05 74.09±0.09 40.90±0.09 2.74±0.04 F test ** ** ** ** 注:* P<0.05的显著水平,** P<0.01的极显著水平,ns为不显著,小写字母表示在0.05水平上显著,下同。
Note: * P<0.05 level of significance, ** P<0.01 level of significance, ns-no significant, lowercase letter indicate significant at 0.05 level, The same below.表 3 造林后2、5、7 a时山地木麻黄18个种源树高、胸径及单株材积的均值及其多重比较结果
Table 3. Average values on height, DBH and individual volume with standard error and Duncan Multiple Range Test of 18 C. junghuhniana provenances at 2, 5 and 7 years after planting
种源
Provenance树高Height /m 胸径DBH/cm 单株材积Volume/×10-3m3 2 a 5 a 7 a 2 a 5 a 7 a 2 a 5 a 7 a 18950 4.54±0.21cdef 6.38±0.29cd 9.25±0.64cdef 3.17±0.21g 5.39±0.36cde 6.89±0.58efg 1.51±0.34def 6.14±1.29def 14.35±3.9def 18954 4.45±0.21def 6.55±0.30cd 9.95±0.54bcde 3.32±0.21defg 5.41±0.36cde 7.37±0.50efg 1.66±0.34cdef 6.51±1.33def 17.38±3.30def 18845 4.90±0.22bcde 5.94±0.32def 8.34±0.57efg 3.76±0.21cdef 5.17±0.39de 6.36±0.53fg 2.59±0.34b 4.96±1.42f 10.44±3.49f 18844 5.05±0.21bc 6.64±0.30cd 8.51±0.51defg 4.09±0.20bc 6.00±0.36bcd 6.59±0.47efg 2.95±0.33b 7.63±1.33cdef 11.14±3.08ef 18846 4.58±0.21cde 6.18±0.29de 8.15±0.49fg 3.34±0.20defg 4.84±0.35e 5.96±0.45g 1.75±0.33cdef 4.98±1.26f 10.20±2.99f 18849 5.03±0.21bcd 7.68±0.30ab 10.82±0.50abc 3.22±0.21g 6.35±0.36bc 7.84±0.46def 1.65±0.33cdef 9.45±1.33cde 19.01±3.04cdef 18850 5.32±0.22b 7.21±0.35bc 10.06±0.57bcd 3.84±0.22cd 6.73±0.42ab 8.14±0.52abcde 2.70±0.35b 9.75±1.52cde 18.47±3.49def 18852 4.64±0.22cde 6.35±0.32cd 8.81±0.55defg 3.27±0.22efg 5.45±0.39cde 6.96±0.50efg 1.73±0.35cdef 5.99±1.42ef 13.64±3.36def 19489 5.45±0.21b 7.92±0.28ab 11.46±0.49ab 4.20±0.21bc 7.59±0.34a 9.18±0.45abcd 2.91±0.33b 14.26±1.25ab 28.33±2.99abc 19490 6.12±0.21a 7.91±0.32ab 11.67±0.55a 4.75±0.20a 7.66±0.38a 9.55±0.51a 4.46±0.33a 15.86±1.38a 34.00±3.36a 17878 4.36±0.21ef 6.61±0.29cd 9.66±0.51cdef 3.36±0.20defg 6.42±0.35bc 8.01±0.46bcde 1.77±0.33cdef 10.03±1.26cde 22.97±3.13bcd 17877 5.96±0.22a 8.26±0.29a 11.45±0.53ab 4.44±0.21ab 7.73±0.36a 9.40±0.48abc 3.87±0.34a 16.14±1.29a 34.88±3.24a 19239 4.46±0.23def 6.58±0.34cd 9.00±0.55defg 3.79±0.23cde 6.46±0.41bc 7.65±0.51ef 2.42±0.36bc 10.10±1.49cd 20.87±3.36cde 19238 4.50±0.23cdef 6.74±0.29cd 10.17±0.58abcd 3.24±0.22fg 6.40±0.36bc 8.96±0.54abcd 1.85±0.36cde 11.43±1.29bc 31.45±3.56ab 19242 4.36±0.21ef 4.98±0.33g 7.61±0.66g 3.40±0.21defg 4.89±0.40de 6.72±0.61efg 2.30±0.34bcd 4.43±1.44f 12.67±4.00def 19241 3.48±0.23g 5.16±0.35fg 8.13±0.64fg 2.66±0.22h 5.13±0.42de 7.18±0.59efg 1.02±0.36f 5.94±1.54ef 19.35±3.90cdef 19237 3.95±0.21fg 5.47±0.31efg 7.50±0.66g 2.96±0.21gh 5.16±0.37de 7.25±0.59efg 1.23±0.33ef 4.87±1.36f 13.51±4.00def 18853 4.40±0.23ef 6.50±0.35cd 9.38±0.64cdef 3.19±0.23g 5.64±0.42bcde 7.36±0.59efg 1.85±0.36cde 7.71±1.52cdef 18.12±3.90def 均值Means 4.77±0.06 6.66±0.08 9.55±0.14 3.57±1.46 6.06±0.09 7.67±0.13 2.25±0.09 8.82±0.35 19.76±0.87 F test ** ** ** ** ** ** ** ** ** 表 4 造林后7 a时18个种源山地木麻黄形质性状的得分均值及其多重比较结果
Table 4. The mean score of qualitative traits with standard error and Duncan Multiple Range Test of 18 C. junghuhniana provenances at 7years after planting
种源
ProvenanceRES DPB TPB APB LPB LDB AP SFS 18950 3.13±0.06abc 3.33±0.05abc 3.25±0.08a 1.42±0.06bcd 1.92±0.03a 1.33±0.06bcd 4.79±0.08abcd 2.82±0.03efg 18954 3.40±0.05abc 3.06±0.05bc 3.00±0.07abcd 1.38±0.05bcd 1.94±0.03a 1.56±0.05abc 4.61±0.08abcd 3.29±0.03ab 18845 3.46±0.06ab 3.22±0.06bc 3.11±0.09ab 1.22±0.07d 2.00±0.04a 1.67±0.07ab 4.00±0.09cd 2.84±0.04efg 18844 3.24±0.05abc 3.45±0.04ab 3.05±0.06abcd 1.32±0.04cd 1.95±0.03a 1.64±0.04ab 4.58±0.08abcd 2.89±0.04def 18846 3.40±0.05ab 3.44±0.05ab 3.06±0.07abc 1.19±0.05d 2.00±0.03a 1.50±0.05abc 4.16±0.08bcd 3.20±0.03bcd 18849 3.60±0.05a 3.20±0.04bc 2.70±0.06abcde 1.30±0.05cd 2.00±0.03a 1.35±0.05bcd 4.21±0.08bcd 3.56±0.03a 18850 3.13±0.05abc 3.06±0.05bc 2.44±0.07bcde 1.44±0.05abcd 1.94±0.03a 1.44±0.05abc 3.85±0.09d 3.09±0.04bcde 18852 3.31±0.05abc 3.26±0.04bc 2.95±0.06abcd 1.47±0.05abcd 1.89±0.03a 1.32±0.05bcd 4.39±0.08abcd 3.08±0.04bcde 19489 3.27±0.05abc 3.43±0.04ab 2.62±0.06abcde 1.71±0.05ab 1.95±0.03a 1.67±0.04ab 4.91±0.08abc 3.20±0.03bcd 19490 3.25±0.05abc 3.31±0.05abc 2.38±0.07de 1.50±0.05abcd 2.00±0.03a 1.69±0.05ab 4.49±0.08abcd 3.03±0.04bcde 17878 3.00±0.05bc 3.29±0.04abc 2.43±0.06cde 1.81±0.05a 1.86±0.03a 1.00±0.04d 4.23±0.08abcd 2.63±0.03fg 17877 3.28±0.05abc 3.50±0.04ab 2.70±0.06abcde 1.75±0.05ab 1.95±0.03a 1.80±0.05a 4.95±0.08ab 3.15±0.03bcde 19239 3.30±0.06abc 3.26±0.04bc 2.87±0.06abcde 1.30±0.04cd 2.00±0.03a 1.61±0.04ab 4.75±0.08abcd 3.06±0.04bcde 19238 3.28±0.06abc 3.16±0.04bc 2.32±0.06e 1.47±0.05abcd 1.95±0.03a 1.37±0.05bcd 4.97±0.08ab 3.05±0.03bcde 19242 3.14±0.06abc 3.33±0.05abc 2.67±0.08abcde 1.67±0.06abc 2.00±0.03a 1.00±0.06d 4.34±0.08abcd 2.51±0.04g 19241 3.00±0.06abc 3.44±0.05ab 2.94±0.07abcde 1.38±0.05bcd 2.00±0.03a 1.31±0.05bcd 4.35±0.08abcd 2.89±0.04cdef 19237 2.81±0.06c 3.79±0.05a 3.14±0.07ab 1.14±0.06d 1.86±0.03a 1.43±0.05abc 5.11±0.08a 2.86±0.04def 18853 3.40±0.06abc 3.31±0.05c 2.69±0.08bcde 1.46±0.06d 1.92±0.03a 1.38±0.06cd 5.00±0.09ab 3.26±0.04abc 均值Means 3.25±0.01 3.32±0.01 2.77±0.02 1.45±0.01 1.95±0.01 1.46±0.01 4.54±0.02 3.03±0.01 F test ns ns ** ** ns ** * ** 表 5 山地木麻黄各性状的方差分量(种源、互作、表型和误差)、种源遗传力、遗传变异系数及遗传增益
Table 5. Estimates of variances (σp2, σp2R, σph2, σe2) for additive, interaction, phenotypic and error, respectively, and Provenance heritability (HP2), genetic coefficient of variation (GCVA) and genetic gains (ΔG) for various traits
性状Trait σp2 σp2R σph2 σp2e Hp2/% GCVA/% ΔG/% H2 0.344 0.581 2.339 1.414 14.70** 12.30 2.92 DBH2 0.182 0.767 2.104 1.155 8.64* 11.96 1.95 V2 0.461a 2.590a 5.481a 2.430a 8.42* 30.20 4.86 H5 0.666 0.648 4.164 2.850 16.00** 12.25 3.08 DBH5 0.685 0.846 5.778 4.247 11.90** 13.67 2.69 V5 0.011b 0.012b 0.079b 0.055b 14.30** 38.11 9.08 H7 1.290 1.550 9.370 6.530 13.80** 11.89 2.59 DBH7 0.802 1.199 7.734 5.733 10.40* 11.67 2.17 V7 0.047b 0.061b 0.355b 0.247b 13.40** 34.85 8.41 RES 0.439b 0.002 0.080 0.077 0.50ns 0.64 0.03 DPB 0.016a 0.003 0.033 0.030 0.05a** 3.84a 3.29b TPB 0.005 0.003 0.077 0.069 6.88* 2.62 0.91 APB 0.004 0.003 0.047 0.040 8.75* 4.41 1.70 LPB 0.043b 0.002 0.014 0.012 0.30 ns 0.34 0.01 LDB 0.007 0.001 0.046 0.039 14.30** 5.56 2.45 AP 0.002 0.019 0.248 0.227 0.72 ns 0.93 0.07 SFS 0.003 0.009 0.051 0.038 6.54* 1.91 0.65 注:表中a表示方差分量×10-3, b表示方差分量×10-6;H2、DBH2、V2、H5、DBH5、V5、H7、DBH7、V7分别表示造林后2、5、7 a时的树高、胸径和单株材积;RES、DPB、TPB、APB、LPB、LDB、AP、SFS为造林后7 a时测定;下同
Note: a represents variances×10-3, b represents variances×10-6. H2, DBH2, V2, H5, DBH5, V5, H7, DBH7, V7 represents the height, diameter at breast height and volume at 2, 5 and 7 years after planting, respectively. RES, DPB, TPB, APB, LPB, LDB, AP, SFS is measured at 7 years after planting. The same below.表 6 山地木麻黄各性状的遗传相关系数(上三角),表型相关系数(下三角)
Table 6. Genotypic (upper triangle), phenotypic correlations (lower triangle) for various traits
H2 H5 H7 DBH2 DBH5 DBH7 V2 V5 V7 TPB APB LDB AP SFS H2 0.97** 0.94** 0.94** 0.87** 0.82** 0.92** 0.86** 0.75** -0.52* 0.45* 0.77** -0.17ns 0.38* H5 0.83** 0.99** 0.84** 0.91** 0.85** 0.84** 0.89** 0.78** -0.63** 0.51* 0.69** 0.10ns 0.66** H7 0.77** 0.87** 0.82** 0.78** 0.89** 0.81** 0.74** 0.94** -0.76** 0.67** 0.50* 0.26ns 0.60** DBH2 0.88** 0.74** 0.68** 0.88** 0.84** 0.87** 0.93** 0.80** -0.52* 0.53* 0.86** -0.02ns 0.11ns DBH5 0.77** 0.86** 0.94** 0.81** 0.93** 0.90** 0.94** 0.85** -0.90** 0.76** 0.62** 0.37ns 0.35ns DBH7 0.75** 0.83** 0.82** 0.77** 0.99** 0.85** 0.89** 0.98** -0.92** 0.81** 0.42* 0.75* 0.28ns V2 0.86** 0.76** 0.68** 0.99** 0.82** 0.77** 0.91** 0.84** -0.54* 0.51* 0.80** 0.01ns 0.06ns V5 0.72** 0.85** 0.74** 0.74** 0.99** 0.99** 0.82** 0.89** -0.84** 0.69** 0.61** 0.47ns 0.36ns V7 0.70** 0.79** 0.72** 0.72** 0.96** 0.94** 0.77** 0.99** -0.88** 0.75** 0.40ns 0.75* 0.25ns TPB -0.43** -0.50** -0.55** -0.52* -0.56** -0.60** -0.47** -0.48** -0.55** -0.82** 0.15ns -0.06ns -0.07ns APB 0.23** 0.28** 0.30** 0.23** 0.30** 0.29** 0.20** 0.24** 0.25** -0.15* -0.25ns 0.11ns -0.42ns LDB 0.15* 0.11* 0.09ns 0.16* 0.03ns -0.02ns 0.15* 0.07ns 0.04ns 0.27** - 0.35ns 0.55* AP 0.04ns 0.04ns 0.05ns 0.06* -0.04ns -0.03ns 0.06ns - 0.03ns -0.07ns 0.01ns 0.07ns -0.16ns SFS 0.15** 0.20** 0.17** 0.05ns 0.07* 0.05ns 0.07ns 0.11* 0.10* -0.10* -0.09ns 0.07ns 0.25** 表 7 参试18个种源综合排名
Table 7. Ordinal ranking of 18 C. junghuhniana provenances
种源
ProvenanceH7 D7 V7 SUR TPB APB LDB AP SFS 综合评价
Overview排名
Rank18950 0.042 9 0.077 9 0.346 5 0.265 2 0.000 0 0.047 1 0.067 2 0.000 0 0.043 5 0.890 3 14 18954 0.021 7 0.052 1 0.251 7 0.025 3 0.005 9 0.057 7 0.017 4 0.001 5 0.005 8 0.439 1 5 18845 0.081 3 0.111 9 0.491 1 0.039 4 0.001 8 0.105 3 0.005 5 0.027 5 0.040 8 0.904 7 15 18844 0.073 2 0.095 9 0.463 2 0.002 0 0.004 0 0.073 7 0.008 3 0.011 0 0.035 3 0.766 5 13 18846 0.091 0 0.141 7 0.500 5 0.000 0 0.003 3 0.118 2 0.027 8 0.034 9 0.010 2 0.927 5 16 18849 0.005 3 0.032 1 0.207 1 0.002 8 0.028 6 0.079 3 0.062 5 0.031 6 0.000 0 0.449 3 6 18850 0.019 0 0.021 9 0.221 4 0.108 4 0.062 5 0.042 3 0.040 6 0.061 3 0.017 6 0.594 8 9 18852 0.059 8 0.073 4 0.370 9 0.052 7 0.008 7 0.034 4 0.072 4 0.020 1 0.018 2 0.710 6 10 19489 0.000 3 0.001 5 0.035 2 0.000 6 0.037 7 0.002 8 0.005 5 0.001 6 0.010 2 0.095 4 2 19490 0.000 0 0.000 0 0.000 6 0.068 1 0.072 5 0.029 3 0.003 9 0.015 1 0.022 7 0.212 2 3 17878 0.029 5 0.026 2 0.116 5 0.008 7 0.063 9 0.000 0 0.197 5 0.030 2 0.069 4 0.542 0 8 17877 0.000 4 0.000 2 0.000 0 0.029 5 0.028 6 0.001 1 0.000 0 0.001 0 0.013 7 0.074 5 1 19239 0.052 2 0.039 7 0.161 3 0.085 6 0.013 7 0.077 9 0.011 3 0.005 1 0.020 4 0.467 3 7 19238 0.016 5 0.003 9 0.009 7 0.154 2 0.082 6 0.034 4 0.057 5 0.000 8 0.020 9 0.380 5 4 19242 0.121 2 0.087 8 0.405 4 0.317 0 0.032 2 0.006 2 0.197 5 0.022 8 0.086 8 1.277 0 18 19241 0.092 2 0.061 7 0.198 1 0.209 9 0.009 2 0.057 7 0.073 4 0.022 3 0.035 6 0.760 0 11 19237 0.127 6 0.058 1 0.375 3 0.252 2 0.001 1 0.135 7 0.042 6 0.000 0 0.039 3 1.031 8 17 18853 0.038 6 0.052 7 0.230 8 0.312 6 0.029 4 0.037 0 0.053 3 0.000 5 0.007 4 0.762 3 12 -
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