台湾桤木无性系苗木快繁与多性状综合选择研究
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摘要
台湾桤木(Alnus formosana)原产于中国台湾,在原产地高达30m以上,胸径30cm以上,以生长快、干形通直、纹理直、易于加工等特点被广泛栽培。它是生产木地板、家俱等的优质木材,也可作刨花板、造纸原料。随着加工业的发展,台湾桤木的木材一直都供不应求,导致台湾桤木无性系苗木供不应求,为了满足生产对台湾桤木无性系苗木的需要,研究台湾桤木无性系快繁技术及开展台湾桤木多性状优良无性系综合选育就显得非常必要。
本研究选用湖南省林业科学院初选出的台湾桤木无性系,采用完全随机区组试验设计方法,研究了台湾恺木无性系苗木快繁与选择、光合生理、营养生长、生殖生长及木材材性以及相互关系,并开展了多性状综合选择研究等。主要结论如下：
1、台湾桤木无性系遗传特性、栽植密度、植株定干高度、修枝、施肥等影响台湾桤木采穗圃无性系植株的萌芽数。选择出了2个高产穗条无性系：湘桤F03、湘桤F04。建立台湾桤木无性系采穗圃,栽植密度以株行距1m×2m为宜,无性系植株定干高度以90cm为宜：。
2、台湾桤木植株不同年龄其枝条扦插生根率有显著差异,4年生以上植株扦插生根率显著降低。不同无性系扦插生根率有显著差异,选择出了扦插生根率达80%以上的15个无性系即湘恺F01、湘桤F02、湘桤F03、湘恺F04、洲桤F05、湘桤F06、湘桤F07、湘桤F08、湘桤F09、、湘桤F10、湘桤F11、湘桤F12、湘桤F13、湘桤F14和湘桤F15。在简易条件下,扦插时间以9月上旬扦插生根率最高。扦插基质以纯沙土扦插成活率最高。穗条带顶芽或带1～2片叶可显著提高穗条生根率。不同浓度GGR生根剂处理后的穗条生根率有显著差异,以300mg/L浓度处理穗条切口其扦插生根率最佳。
3、台湾桤木插条不定根形成有皮部生根和愈伤组织生根两种生根类型,大多数无性系呈现皮部生根类型,少数无性系同时包含两种生根类型；台湾桤木无性系生根能力与生根率之间成正相关关系。
4、建立了台湾桤木苗木培育单位而积苗木株数与苗高生长、地径生长关系的数学模型：单位面积苗木数量与苗木平均高度的数学模型为y=2.0147x-30.9568(50≤x≤70)；单位面积苗木数量与苗木平均地径的数学模型为y156.3578-169.1596x(0.4≤x≤0.6)。
5、光合生理研究表明：台湾桤木光饱和点、补偿点及表观量子效率分别为1177.8μmol·m-2·s-1、20.0μmol·m-2·s-1、0.0345,说明台湾桤木利用弱光的能力比较强。
6、氮、磷、钾肥对台湾桤木的光合特性有一定的影响,在25个处理中,处理N12P12K9光合能力最强。光合速率与蒸腾速率、光合有效辐射、气温和叶温呈显著正相关,胞间CO2浓度与光合速率、蒸腾速率呈显著负相关关系,叶片水压亏缺受相对空气湿度的影响大,空气相对湿度受气温、光强影响显著,与蒸腾速率呈显著负相关。
7、选育出了4个生长性状优良的台湾桤木无性系：F02、F07、F12和F21。
8、参试无性系木材基本密度为0.4747g.cm-3～0.3032g.cm-3,选育出了3个木材基本密度较大的台湾桤木无性系：F02、F12和F15。
9、参试无性系木材纤维素含量为45%～55%,选育出了4个木材纤维素含量较大的台湾桤木无性系：F02、F11、F12和F15。
10、台湾桤木木材纤维长度属于中级长度,纤维长宽比均大于35,其纤维长宽比的变动幅度主要集中在37～50之间,纤维长度、宽度和长宽比在数值组间变化差异小,在无性系间差异不显著。
11、9a生台湾桤木基本密度变异属于从髓心向外以曲线形式缓慢增加的模式，且年轮段间差异水平不显著。参试的台湾桤木5个无性系纤维素含量在年轮段间的差异较大。纤维长度的径向变异总体仍为上升趋势,纤维宽度在年轮段间的差异较大,长宽比随着生长轮数的增加也随之增加,1-3年轮段到4-6年轮段的增幅为5.49%,4-6年轮段到7-9年轮段的增幅为3.21%。
12、采用生长作为选择的主程序,而将材性改良作为次程序的育种路线,对台湾桤木无性系多性状进行选择,在国内首次选出了3个生长和材性兼优的台湾桤木无性系,即：F02、F07、F12。
13、参试台湾桤木10个无性系种子千粒重差别较大,种子千粒重最大的是无性系F05,其种子千粒重达到了1.09g,种子千粒重最小的是无性系F09,只有0.42g,单果出籽率的变异系数(CV)的变异幅度为25.00%～37.42%。球果长、宽和鲜重在无性系间都存在显著差异,综合分析无性系F05是参试10个无性系中种实性状最优且最稳定。
Taiwan alder (Alnus formosana) originated in China Taiwan was up to more than30m in height and to more than30cm in DBH in the origin. It has been widely cultivated due to good characteristics of fast growth, straight trunk form, softy timber, straight texture, easy processing. It has been used as high-quality materials for making wood floor and furniture, also for making particleboard and paper making. With the industrial development, Taiwan alder seedlings has been in short supply, the market demand of Taiwan alder seedlings is growing. In order to meet the needs of production on clones seedlings of Taiwan Alder, rapid propagation techniques of clones and comprehensive breeding of multiple characters of excellent clones in Taiwan Alders have been necessarily studied.
Based on clones varieties of Taiwan Alder selected by Hunan Academy of Forestry, rapid propagation and selection, photosynthesis, vegetative growth, reproductive growth and wood properties of clones seedlings of Taiwan alder had been studied by the experiment method of completely randomized block design. The main conclusions were showed as follows:
1. Bud numbers of Taiwan alder clone of cutting orchard depended mainly on the genetic characteristics and planting density, but also closely related with the stem height, pruning, fertilization.2kinds of high-yielding clones cuttings, Xiang Qi F03and Xiang Qi F04, were selected from Taiwan alder clones.1m×2m of planting density and90cm of plant stem height were appropriate for the establishment of the Taiwan alder clones.
2. Cuttings rooting rate of Taiwan alder showed significantly difference among clones of different ages. A significant reduction in cuttings rooting rate were showed in more than4years old.15kinds of clone varieties which rooting rate reached more than80%were selected, they were Xiangqi F01, Xiangqi F02, Xiangqi F03, Xiangqi F04, Xiangqi F05, Xiangqi F06, Xiangqi F07, Xiangqi F08, Xiangqi F09,Xiangqi F10, Xiangqi F11, Xiangqi F12, Xiangqi F13, Xiangqi F14and Xiangqi F15. In simple conditions, early September was the best as cutting time because of the highest rooting rate, pure sand was the best as cutting medium because of the highest survival rate. Cutting with terminal bud or1to2leaves significantly increased the rooting rate. There were significant difference of rooting rate among cutting treated by different concentrations GGR, the effect of300mg/L concentration GGR was the best.
3. Adventitious root of Taiwan alder cuttings formed two types of thedermal rooting and callus rooting, the most of cutting roots belonged to thedermal rooting, but few of cutting roots contained these two types. The rooting ability of Taiwan alder clones were positive related with the rooting rates.
4. The mathematical models were established between the seedling numbers per unit area and the height growth or diameter growth of Taiwan alder, respectively. The relationship model of the numbers per unit area and the average height of seedlings were y=2.0147x-30.9568(50≤x≤70). The relationship model of the numbers per unit area and the average diameter of seedlings were y=156.3578—169.1596x (0.4 5. The researches of photosynthetic physiology were showed that light saturation point, the compensation point and apparent quantum efficiency of Taiwan alder were1177.8μmol·m-2·s-1,20.0μmol·m-2·s-1,0.0345, respectively. It illustrated Taiwan alder could enough used low light.
6. The impact of Nitrogen, phosphorus, potash fertilizer on photosynthetic characteristics of Taiwan alder was observed. Photosynthesis of Taiwan alder treated with N12P12K9fertilizer was strongest in25kinds of fertilizer treatment. The Photosynthetic rate was positively related with transpiration rate, photosynthetic active radiation, air temperature and leaf temperature, respectively. Intercellular CO2concentration was negatively with photosynthetic rate and transpiration rate, respectively. Leaf water pressure deficit was influenced by air relative humidity. Air relative humidity obviously affected by temperature, light intensity, and was significantly negatively correlated with transpiration rate.
7. Bred four excellent growth traits alder clones:4clones varieties of Taiwan alder were selected as excellent growth traits. They were F02, F07, F12and F21.
8. The range of basic wood density from the clones tested were0.4747g.cm-30.3032g.cm-3. the3clones varieties of the larger basic wood density were selected, they were F02、F12and F15.
9. The range of wood cellulose content from the clones tested were45%to55%, the3clones varieties of the larger cellulose content were selected, they were F02, F11, F12and F15.
10. The wood fiber length of Taiwan alder belonged to intermediate length. Fiber length-width ratio were greater than35, and the range of fiber length-width ratio were mainly between37～50. The difference of the fiber length, width and length-width ratio were small in arithmetic groups. The difference of them was not significant between the clones.
11. Density variation of9years old Taiwan alder belonged to the model which was slowly increased from the pith outward curve, and no significant difference of density was observed between annual ring segments. Cellulose contents of5clones of Taiwan alder tested were different at annual ring segments. Radial variation of fiber length was still an upward trend, fiber width were different largely between annual ring segments. The ratio of fiber length to width increased with increasing in the growth rings, the ratio in4-6years of growth rings increased by5.49%over1-3years of growth rings, the ratio in7-9years of growth rings increased by3.21%over4-6years of growth rings.
12. Using breeding model in which the growth was made as the main program of clones selection and the improvement of wood properties were made as the sub program of clones selection, multiple traits of Taiwan alder clones were selected.3clones of Taiwan alder which the best in the growth and wood property were firstly selected, they were F02, F07, F12.
13. Seed grain weights of10clones were largely different. The seed grain weight of clones F05which were up to1.09g was maximum, the seed grain weight of clones F05which were up to0.42g was minimum. The coefficient of variation (CV) of single fruit seed rate ranged from25%to37.42%. There were significant difference of the cone length, width and fresh weight between the clones tested. A comprehensive analysis were done that the clones F05was the best in the seed traits among all of the clones tested in our study.
本研究选用湖南省林业科学院初选出的台湾桤木无性系,采用完全随机区组试验设计方法,研究了台湾恺木无性系苗木快繁与选择、光合生理、营养生长、生殖生长及木材材性以及相互关系,并开展了多性状综合选择研究等。主要结论如下：
1、台湾桤木无性系遗传特性、栽植密度、植株定干高度、修枝、施肥等影响台湾桤木采穗圃无性系植株的萌芽数。选择出了2个高产穗条无性系：湘桤F03、湘桤F04。建立台湾桤木无性系采穗圃,栽植密度以株行距1m×2m为宜,无性系植株定干高度以90cm为宜：。
2、台湾桤木植株不同年龄其枝条扦插生根率有显著差异,4年生以上植株扦插生根率显著降低。不同无性系扦插生根率有显著差异,选择出了扦插生根率达80%以上的15个无性系即湘恺F01、湘桤F02、湘桤F03、湘恺F04、洲桤F05、湘桤F06、湘桤F07、湘桤F08、湘桤F09、、湘桤F10、湘桤F11、湘桤F12、湘桤F13、湘桤F14和湘桤F15。在简易条件下,扦插时间以9月上旬扦插生根率最高。扦插基质以纯沙土扦插成活率最高。穗条带顶芽或带1～2片叶可显著提高穗条生根率。不同浓度GGR生根剂处理后的穗条生根率有显著差异,以300mg/L浓度处理穗条切口其扦插生根率最佳。
3、台湾桤木插条不定根形成有皮部生根和愈伤组织生根两种生根类型,大多数无性系呈现皮部生根类型,少数无性系同时包含两种生根类型；台湾桤木无性系生根能力与生根率之间成正相关关系。
4、建立了台湾桤木苗木培育单位而积苗木株数与苗高生长、地径生长关系的数学模型：单位面积苗木数量与苗木平均高度的数学模型为y=2.0147x-30.9568(50≤x≤70)；单位面积苗木数量与苗木平均地径的数学模型为y156.3578-169.1596x(0.4≤x≤0.6)。
5、光合生理研究表明：台湾桤木光饱和点、补偿点及表观量子效率分别为1177.8μmol·m-2·s-1、20.0μmol·m-2·s-1、0.0345,说明台湾桤木利用弱光的能力比较强。
6、氮、磷、钾肥对台湾桤木的光合特性有一定的影响,在25个处理中,处理N12P12K9光合能力最强。光合速率与蒸腾速率、光合有效辐射、气温和叶温呈显著正相关,胞间CO2浓度与光合速率、蒸腾速率呈显著负相关关系,叶片水压亏缺受相对空气湿度的影响大,空气相对湿度受气温、光强影响显著,与蒸腾速率呈显著负相关。
7、选育出了4个生长性状优良的台湾桤木无性系：F02、F07、F12和F21。
8、参试无性系木材基本密度为0.4747g.cm-3～0.3032g.cm-3,选育出了3个木材基本密度较大的台湾桤木无性系：F02、F12和F15。
9、参试无性系木材纤维素含量为45%～55%,选育出了4个木材纤维素含量较大的台湾桤木无性系：F02、F11、F12和F15。
10、台湾桤木木材纤维长度属于中级长度,纤维长宽比均大于35,其纤维长宽比的变动幅度主要集中在37～50之间,纤维长度、宽度和长宽比在数值组间变化差异小,在无性系间差异不显著。
11、9a生台湾桤木基本密度变异属于从髓心向外以曲线形式缓慢增加的模式，且年轮段间差异水平不显著。参试的台湾桤木5个无性系纤维素含量在年轮段间的差异较大。纤维长度的径向变异总体仍为上升趋势,纤维宽度在年轮段间的差异较大,长宽比随着生长轮数的增加也随之增加,1-3年轮段到4-6年轮段的增幅为5.49%,4-6年轮段到7-9年轮段的增幅为3.21%。
12、采用生长作为选择的主程序,而将材性改良作为次程序的育种路线,对台湾桤木无性系多性状进行选择,在国内首次选出了3个生长和材性兼优的台湾桤木无性系,即：F02、F07、F12。
13、参试台湾桤木10个无性系种子千粒重差别较大,种子千粒重最大的是无性系F05,其种子千粒重达到了1.09g,种子千粒重最小的是无性系F09,只有0.42g,单果出籽率的变异系数(CV)的变异幅度为25.00%～37.42%。球果长、宽和鲜重在无性系间都存在显著差异,综合分析无性系F05是参试10个无性系中种实性状最优且最稳定。
Taiwan alder (Alnus formosana) originated in China Taiwan was up to more than30m in height and to more than30cm in DBH in the origin. It has been widely cultivated due to good characteristics of fast growth, straight trunk form, softy timber, straight texture, easy processing. It has been used as high-quality materials for making wood floor and furniture, also for making particleboard and paper making. With the industrial development, Taiwan alder seedlings has been in short supply, the market demand of Taiwan alder seedlings is growing. In order to meet the needs of production on clones seedlings of Taiwan Alder, rapid propagation techniques of clones and comprehensive breeding of multiple characters of excellent clones in Taiwan Alders have been necessarily studied.
Based on clones varieties of Taiwan Alder selected by Hunan Academy of Forestry, rapid propagation and selection, photosynthesis, vegetative growth, reproductive growth and wood properties of clones seedlings of Taiwan alder had been studied by the experiment method of completely randomized block design. The main conclusions were showed as follows:
1. Bud numbers of Taiwan alder clone of cutting orchard depended mainly on the genetic characteristics and planting density, but also closely related with the stem height, pruning, fertilization.2kinds of high-yielding clones cuttings, Xiang Qi F03and Xiang Qi F04, were selected from Taiwan alder clones.1m×2m of planting density and90cm of plant stem height were appropriate for the establishment of the Taiwan alder clones.
2. Cuttings rooting rate of Taiwan alder showed significantly difference among clones of different ages. A significant reduction in cuttings rooting rate were showed in more than4years old.15kinds of clone varieties which rooting rate reached more than80%were selected, they were Xiangqi F01, Xiangqi F02, Xiangqi F03, Xiangqi F04, Xiangqi F05, Xiangqi F06, Xiangqi F07, Xiangqi F08, Xiangqi F09,Xiangqi F10, Xiangqi F11, Xiangqi F12, Xiangqi F13, Xiangqi F14and Xiangqi F15. In simple conditions, early September was the best as cutting time because of the highest rooting rate, pure sand was the best as cutting medium because of the highest survival rate. Cutting with terminal bud or1to2leaves significantly increased the rooting rate. There were significant difference of rooting rate among cutting treated by different concentrations GGR, the effect of300mg/L concentration GGR was the best.
3. Adventitious root of Taiwan alder cuttings formed two types of thedermal rooting and callus rooting, the most of cutting roots belonged to thedermal rooting, but few of cutting roots contained these two types. The rooting ability of Taiwan alder clones were positive related with the rooting rates.
4. The mathematical models were established between the seedling numbers per unit area and the height growth or diameter growth of Taiwan alder, respectively. The relationship model of the numbers per unit area and the average height of seedlings were y=2.0147x-30.9568(50≤x≤70). The relationship model of the numbers per unit area and the average diameter of seedlings were y=156.3578—169.1596x (0.4
6. The impact of Nitrogen, phosphorus, potash fertilizer on photosynthetic characteristics of Taiwan alder was observed. Photosynthesis of Taiwan alder treated with N12P12K9fertilizer was strongest in25kinds of fertilizer treatment. The Photosynthetic rate was positively related with transpiration rate, photosynthetic active radiation, air temperature and leaf temperature, respectively. Intercellular CO2concentration was negatively with photosynthetic rate and transpiration rate, respectively. Leaf water pressure deficit was influenced by air relative humidity. Air relative humidity obviously affected by temperature, light intensity, and was significantly negatively correlated with transpiration rate.
7. Bred four excellent growth traits alder clones:4clones varieties of Taiwan alder were selected as excellent growth traits. They were F02, F07, F12and F21.
8. The range of basic wood density from the clones tested were0.4747g.cm-30.3032g.cm-3. the3clones varieties of the larger basic wood density were selected, they were F02、F12and F15.
9. The range of wood cellulose content from the clones tested were45%to55%, the3clones varieties of the larger cellulose content were selected, they were F02, F11, F12and F15.
10. The wood fiber length of Taiwan alder belonged to intermediate length. Fiber length-width ratio were greater than35, and the range of fiber length-width ratio were mainly between37～50. The difference of the fiber length, width and length-width ratio were small in arithmetic groups. The difference of them was not significant between the clones.
11. Density variation of9years old Taiwan alder belonged to the model which was slowly increased from the pith outward curve, and no significant difference of density was observed between annual ring segments. Cellulose contents of5clones of Taiwan alder tested were different at annual ring segments. Radial variation of fiber length was still an upward trend, fiber width were different largely between annual ring segments. The ratio of fiber length to width increased with increasing in the growth rings, the ratio in4-6years of growth rings increased by5.49%over1-3years of growth rings, the ratio in7-9years of growth rings increased by3.21%over4-6years of growth rings.
12. Using breeding model in which the growth was made as the main program of clones selection and the improvement of wood properties were made as the sub program of clones selection, multiple traits of Taiwan alder clones were selected.3clones of Taiwan alder which the best in the growth and wood property were firstly selected, they were F02, F07, F12.
13. Seed grain weights of10clones were largely different. The seed grain weight of clones F05which were up to1.09g was maximum, the seed grain weight of clones F05which were up to0.42g was minimum. The coefficient of variation (CV) of single fruit seed rate ranged from25%to37.42%. There were significant difference of the cone length, width and fresh weight between the clones tested. A comprehensive analysis were done that the clones F05was the best in the seed traits among all of the clones tested in our study.
引文
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