Effects of artificial promotion measures on seed germination and early growth of Larix principis-rupprechtii
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摘要:目的 为了解决塞罕坝地区华北落叶松人工林林下更新困难的问题，实现人工林近自然经营，开展人工促进措施（以下简称促更）对林下更新效果的研究十分必要。方法 以塞罕坝地区31年生华北落叶松人工林为研究对象，采用裂区设计等试验方法，从林分密度、枯落物处理方式、种子来源以及种子处理方式这4个方面进行对比试验，探讨林下更新对不同促更措施的响应。结果 处理后的华北落叶松林下均能更新，更新频率为100%左右。同时期不同密度的幼苗地径、苗高在生长末期的差异较小（P > 0.05），而幼苗地径在2年间存在显著性差异（P < 0.05）。不同处理间生长季成活率无显著差异（P > 0.05），整体维持在43.68% ~ 87.87%范围内；而更新密度、播种发芽率在不同枯落物处理方式间差异显著（P < 0.05）。人工播种时，以沟状清理（SC）的更新密度最大，为8 633.33 株/hm2，该处理下种子发芽率最高，为1.79%。此外，更新动态因播种时间而发生改变。自然下种时，处理间的更新动态相对一致，7月底为幼苗萌发高峰期；6月中旬人工播种时，只有SC处理的种子在7月底适时大量萌发。结论 清理枯落物层和播种对于促进华北落叶松林下更新是有效的。促更措施主要是通过影响幼苗数量而非幼苗品质来影响更新效果。播种时间和种子处理方式会对幼苗的更新动态产生影响，6月中旬人工播种无处理的种子会造成当年更新动态的延迟。本试验条件下最佳促更措施：一是在枯落物沟状清理后6月中旬前进行人工播种；二是全面清理枯落物，同时在前一年度的11月初人工播种。Abstract:Objective In order to solve the problem of understory regeneration of Larix principis-rupprechtii plantation in Saihanba Area of Hebei Province, northern China and realize the near-natural management of plantation, it is necessary to study the effects of artificial promotion measures (hereinafter referred to as “promotion”) on understory regeneration.Method This study took 31 years old L. principis-rupprechtii plantation in Saihanba Area as the research object, we adopted the split plot design and other methods. The experiments of promotion were carried out from four aspects: stand density, litter treatment, seed source and seed treatment. This paper discusses the response of understory regeneration to different promotion measures.Result The results showed that the regeneration can appear among the treatments and the regeneration frequency was about 100%. The ground diameter and seedling height of seedlings with different densities at the same period had little difference at the end of growth (P > 0.05), while the ground diameter of seedlings had significant differences in two years (P < 0.05). There was no significant difference in survival rate between different treatments (P >0.05), and the overall survival rate remained in the range of 43.68%−87.87%. However, the regeneration density and germination rate of seeds were significantly different among litter treatments (P < 0.05). The regeneration density of 8 633.33 plant/ha of sulcate cleaning (SC) was the highest under artificial sowing, and seed germination rate of 1.79% was also the highest. In addition, the dynamic of regeneration changed with sowing time. When the seeds were sown naturally, the regeneration dynamics among the treatments were relatively consistent, and the peak of seedling germination was at the end of July. When the seeds were sown artificially in the middle of June, only the seeds in SC treatment germinated in large quantities at the end of July.Conclusion The artificial interference with litter layer and seeding are effective to promote the regeneration of L. principis-rupprechtii plantations. The promotion measures mainly affect the effects of regeneration by affecting the number of seedlings rather than the seedling quality. The sowing time and seed treatments affect the renewal dynamics of seedlings, and artificial sowing of untreated seeds in mid-June can cause the delay of renewal dynamics in the current years. The best way to promote the regeneration of L. principis-rupprechtii plantations is as follows: the first way is to sulcate clean the litter layer and spread the seeds manually before mid-June, the second way is comprehensively clean the litter layer and spread the seeds in early November of the previous year.
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图 1 裂区设计示意图
CK. 对照；CC. 全面清理；BC. 带状清理；SC. 沟状清理。图中4类枯落物处理方式在副区中是随机排列的，林分密度由小到大分别表示为T1、T2、T3。下同。CK, control; CC, completely cleaning; BC, banding cleaning ; SC, sulcate cleaning. The order of 4 litter treatments is random in the subplot, and stand densities from small to large are expressed as T1, T2, T3, respectively. The same below.
Figure 1. Schematic diagram of slip plot design
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图 2 带状清理（BC，a）和沟状清理（SC，b）示意图
灰色区域为副区中枯落物的堆积区，白色区域为副区中枯落物的清理区。The gray area is the accumulation area of litter and the white area is the cleaning area of litter in the figure.
Figure 2. Schematic diagram of banding cleaning (a) and sulcate cleaning (b)
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图 3 幼苗地径、苗高在不同时期的比较
不同大写字母表示密度间的差异显著（P < 0.05），不同小写字母表示年份间的差异显著（P < 0.05）。Different capital letters indicate significant difference between varied densities (P < 0.05), and different small letters indicate significant difference between varied years (P < 0.05).
Figure 3. Comparison of seedling ground diameter and seedling height in different periods
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图 4 2019和2020年不同林分密度下不同枯落物处理方式的更新密度
不同大写字母表示枯落物处理方式间的差异显著（P < 0.05），不同小写字母表示密度间的差异显著（P < 0.05）。图6同。Different capital letters indicate the difference between different litter treatment modes (P < 0.05), and different small letters indicate the difference between varied stand densities (P < 0.05). The same as figure 6.
Figure 4. Regenaration density of different litter treatment modes under varied stand densities in 2019 and 2020
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图 5 T1、T2、T3林分密度下2019和2020年不同处理间生长季存活幼苗密度的动态变化
Figure 5. Dynamic changes of survival seedling density in growing season among different treatments under T1, T2 and T3 stand densities in 2019 and 2020
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图 6 人工播种下各项处理中种子发芽率比较
Figure 6. Comparison of seed germination rate of different treatments under artificial sowing
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图 7 2020年不同种子处理方式间生长季存活幼苗密度的动态变化
Figure 7. Dynamic changes of survival seedling density in growing season among different seed treatments in 2020
表 1 固定试验样地基本林分信息
Table 1 Basic stand information of fixed experimental sample plots
样地号
Sample plot No.平均密度/（株ˑhm−2）
Average density/(treeˑha−1)平均胸径
Average DBH/cm平均树高
Average tree height/m平均冠幅
Average crown width/m郁闭度
Canopy density海拔
Altitude/m立地类型
Site type土壤类型
Soil type土层状况
Soil conditionT1 1 650 ± 78 14.76 ± 2.73 13.62 ± 1.58 2.45 ± 0.56 0.52 1 669 曼甸
Plateau灰色森林土
Grey forest soil厚土
Thick soilT2 2 250 ± 125 13.29 ± 3.16 12.64 ± 1.96 2.21 ± 0.71 0.62 T3 2 850 ± 209 12.23 ± 3.57 12.71 ± 2.15 1.90 ± 0.83 0.67 TN 1 375 ± 65 14.92 ± 2.97 12.40 ± 1.76 2.59 ± 0.69 0.49 
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表 2 华北落叶松良种基本参数
Table 2 Basic parameters of high quality seeds of L. principis-rupprechtii
平均粒径
Average grain diameter/mm千粒质量
Thousand-grain mass/g发芽率
Germination rate/%生活力
Seed viability/%1.56 ± 0.14 5.90 ± 0.19 88.89 ± 4.16 97.78 ± 1.57
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表 3 裂区设计的样地处理详情
Table 3 Treatment details of fixed experimental sample plots in slip plot design
林分密度
Stand density枯落物处理方式
Litter treatment method枯落物清理面积
Litter clearing area/m2播种种子密度
Seeding density播种种子质量
Quality of seed propagation播散量
Seeding amount/（g·m−2）播种密度/（粒·m−2）
Seeding density/（cap·m−2）各副区质量
Mass of each
subplot/g各副区粒数
Grain number of
each subplotT1 CK 0 1.50 254.15 150.00 25 415.00 CC 100 150.00 25 415.00 BC 50 75.00 12 707.50 SC 19 28.50 4 828.85 T2 CK 0 1.50 254.15 150.00 25 415.00 CC 100 150.00 25 415.00 BC 50 75.00 12 707.50 SC 19 28.50 4 828.85 T3 CK 0 1.50 254.15 150.00 25 415.00 CC 100 150.00 25 415.00 BC 50 75.00 12 707.50 SC 19 28.50 4 828.85
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表 4 4类种子处理方式
Table 4 4 kinds of seed treatment methods
处理
Treatment播种时间
Sowing time越冬保存条件
Wintering preservation condition雪藏方式
Snow storage mode播种前是否催芽
Whether to promote germination before sowingST1 2019年11月上旬
Early November, 2019塞罕坝室外环境
Outdoor environment of Saihanba Area室外天然雪藏
Natural snow stratification in the outdoor environment否 No ST2 2020年6月中旬
Mid-June, 2020塞罕坝室内环境
Indoor environment of Saihanba Area不雪藏
No snow stratification否 No ST3 2020年6月中旬
Mid-June, 2020塞罕坝室内环境
Indoor environment of Saihanba Area不雪藏
No snow stratification是 Yes ST4 2020年6月中旬
Mid-June, 2020塞罕坝室内环境
Indoor environment of Saihanba Area室内人工雪藏
Artificial snow storage in the indoor environment是 Yes
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表 5 2019年和2020年华北落叶松林各级幼苗的更新频率
Table 5 Regeneration frequency of seedlings at various levels in L. principis-rupprechtii plantations in 2019 and 2020
林分
密度
Stand
density枯落物
处理方式
Litter treatment mode2019 2020 1a 更新频率
Regeneration
frequency/%2a 更新频率
Regeneration
frequency/%3a+ 更新频率
Regeneration
frequency/%整体更新频率 Overall
regeneration
frequency/%1a 更新频率
Regeneration
frequency/%2a 更新频率
Regeneration
frequency/%3a+ 更新频率 Regeneration
frequency/%整体更新频率 Overall
regeneration
frequency/%T1 CK √ 100 0 0 100 √ 100 √ 50 0 100 CC √ √ BC √ √ √ SC √ √ T2 CK √ 75 √ 25 0 75 √ 100 √ 25 0 100 CC √ BC √ √ SC √ √ T3 CK √ 100 √ 25 0 100 √ 100 25 √ 25 100 CC √ √ BC √ √ √ SC √ √ 注：1a. 1年生幼苗；2a. 2年生幼苗；3a+. 3年生及以上幼苗。存在更新的样地标注符号“√”，没有更新的样地不作任何标注。幼树更新频率均为0，故在表中省略。Notes: 1a means 1-year-old seedlings; 2a means 2-year-old seedlings; 3a+ means more than 3 years old seedlings. There is a symbol “√” for the regenerated sample plots; there is no mark for the non-regenerated sample plots, and the regeneration frequency of young trees in all sample plots is 0, it is omitted in the above table.
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表 6 2019年和2020年的华北落叶松更新等级评定
Table 6 Evaluation of regeneration levels of L. principis-rupprechtii in 2019 and 2020
林分密度
Stand
density枯落物处理方式
Litter treatment mode2019 2020 更新密度/（株ˑhm−2）
Regeneration density/（treeˑha−1）更新等级
Regeneration grade更新密度/（株ˑhm−2）
Regeneration density/（treeˑha−1）更新等级
Regeneration gradeT1 CK 266.67 ± 47.14a 不良 Bad 1 933.33 ± 758.65a 不良/中等 Bad or medium CC 66.67 ± 94.28b 不良 Bad 5 700.00 ± 1 489.97a 良好 Good BC 300.00 ± 216.02b 不良 Bad 7 466.67 ± 1 901.46a 良好 Good SC 100.00 ± 141.42a 不良 Bad 7 200.00 ± 3 706.75a 中等/良好 Medium or good T2 CK 366.67 ± 449.69b 不良 Bad 1 833.33 ± 205.48a 不良/中等 Bad or medium CC 0.00 ± 0.00b 不良 Bad 4 300.00 ± 565.69a 中等/良好 Medium or good BC 100.00 ± 81.65b 不良 Bad 6 000.00 ± 648.07a 良好 Good SC 100.00 ± 141.42a 不良 Bad 11 100.00 ± 5 114.68a 良好 Good T3 CK 566.67 ± 205.48b 不良 Bad 2 733.33 ± 449.69a 中等 Medium CC 33.33 ± 47.14b 不良 Bad 5 300.00 ± 1 737.81a 中等/良好 Medium or good BC 66.67 ± 94.28a 不良 Bad 6 433.33 ± 3 492.21a 中等/良好 Medium or good SC 100.00 ± 141.42b 不良 Bad 7 600.00 ± 2 698.15a 良好 Good
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表 7 2019年和2020年不同林分密度下幼苗生长季成活率比较
Table 7 Comparison of survival rate of seedlings in growing season under different standdensities in 2019 and 2020
林分密度
Stand density生长季成活率
Survival rate in growing season/%2019 2020 T1 70.28 ± 18.57Aa 73.93 ± 8.54Aa T2 77.22 ± 9.65Aa 62.11 ± 14.97Aa T3 49.65 ± 37.12Aa 62.26 ± 7.67Aa 注：大写字母表示密度间的差异，小写字母表示年份间的差异。字母不同表示差异显著，差异显著性水平0.05。Notes: capital letters indicate the difference between varied densities, and small letters indicate the difference between varied years. Different letters mean significant difference, and the difference significance level is 0.05.
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表 8 2020年不同林分密度和不同处理方式下幼苗生长季成活率比较
Table 8 Comparison of survival rate of seedlings in growing season of different litter treatment modes under varied stand densities in 2020
枯落物处理方式
Litter treatment mode林分密度 Stand density T1 T2 T3 CK 68.25 ± 27.31Aa 57.30 ± 22.44Aa 72.25 ± 10.81Aa CC 65.85 ± 9.16Aa 85.22 ± 16.92Aa 60.50 ± 2.00Aa BC 87.87 ± 1.04Aa 62.22 ± 10.80Aa 65.17 ± 18.27Aa SC 73.74 ± 5.18Aa 43.68 ± 8.25Aa 51.12 ± 16.89Aa 注：大写字母表示枯落物处理方式间的差异，小写字母表示密度间的差异。字母不同表示差异显著，差异显著性水平0.05。下同。Notes: capital letters indicate the difference between varied litter treatments, and small letters indicate the difference between varied stand densities. Different letters mean significant difference, the difference significance level is 0.05. The same below.
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