Why seedlings grow: influence of plant attributes

Successful forest restoration requires planting quality seedlings with optimal growth potential. Thus, nurseries need to produce seedlings with plant attributes that favor the best chance of successful establishment once they are field planted. From the mid-twentieth century on, research foresters have critically examined plant attributes that confer improved seedling growth under various restoration site conditions. This review examines the value of commonly measured seedling quality attributes (i.e., height, diameter, root mass, shoot-to-root ratio, drought resistance, freezing tolerance, nutrient status, root growth potential, and root electrolyte leakage) that have been recognized as important in explaining why seedlings with improved attributes have better growth after planting. Seedlings with plant attributes that fall within the appropriate range of values can increase the speed with which they overcome planting stress, initiate growth, and become “coupled” to the forest restoration site, thereby ensuring successful seedling establishment. Although planting high quality seedlings does not guarantee successful seedling establishment, it increases chances for successful establishment and growth.     

Bareroot versus container stocktypes: a performance comparison

This review examines the published work on bareroot and container stocktypes in forest restoration programs. The objective was to define overall trends between these two stocktypes and describe what they mean in terms of available information on their nursery and field performance. Stock quality assessments show bareroot seedlings have larger shoot systems because they are typically grown at lower densities, and in many instances longer timeframes, than container seedlings. Container systems typically produce seedlings having a lower shoot to root ratio and a greater root growth potential, conferring greater drought avoidance potential. However, assessments of stress resistance and nutrition found no conclusive evidence that either stocktype has a performance advantage, other than the container plug acting as a source of water and nutrient storage available for outplanting performance. Bareroot seedlings are more sensitive to handling practices of lifting, storage, transport and planting and these practices can negatively affect their performance. Container seedlings can have a higher level of field survival which is related, in part, to their greater drought avoidance potential, thereby overcoming planting stress. Bareroot and container seedlings have comparable survival rates on sites with minimal planting stress. Once seedlings are established, bareroot and container seedlings can have comparable field performance. In many instances where plant competition is the main limiting site variable, larger sized bareroot and container stocktypes have the best chance for successful stand establishment. The lack of a natural root form and root distribution for both stocktypes is a debated mechanical stability issue, though risks of windthrow have not been consistently demonstrated for either stocktype.     

Initial seedling morphological characteristics and field performance of two Sudanian savanna species in relation to nursery production period and watering regimes

It has become apparent that some interventions are required to aid the regeneration of woody species in the Sudanian savanna. Direct seeding has been ineffective, thus planting high quality seedlings may be a viable alternative. In this study, we examined the stock quality of two valuable Sudanian savanna species, Acacia macrostachya and Pterocarpus erinaceus. Different nursery production periods were tested as well as the species’ field performance under well-watered and stressed conditions. The results showed that older seedlings (9-month) were morphologically distinct from younger ones (3-month), particularly in the case of P. erinaceus. Eighteen months after planting out, survival and growth of seedlings were not affected by initial seedling size; this was the result of the high root to shoot ratio of seedlings in all age groups at the time of planting. Seedling mortality as high as 30% was observed and attributed to both drought stress and other factors such as herbivory. Regression analyses revealed that initial shoot height was a poor predictor of field performance for both species, but initial root collar diameter accounted for 25% of the variation in diameter of P. erinaceus in the field. We conclude that initial seedling size does not affect survival and growth in the field provided that all sizes of seedling have a high root to shoot ratio at the time of planting. The prediction of field performance could be improved by developing a model that incorporates a wide range of root collar diameter.     

不同育苗方式对薄壳山核桃苗木生长及根系结构的影响

该文报道了薄壳山核桃大田播种育苗和火箭盆、无纺布、塑料钵容器等育苗方式对苗木生长及根系结构的影响。结果表明:不同育苗方式对薄壳山核桃苗木高度和地径生长影响显著,其中火箭盆容器培育的苗高、地径生长量最大,为39.14 cm和0.96 cm;无纺布容器苗和大田播种苗次之;塑料钵容器苗的苗高、地径生长量最小。多重比较分析表明,不同育苗方式对薄壳山核桃苗木的主根长度、主根直径、1级侧根数、1级侧根平均长度、根鲜质量、根干质量、茎鲜质量、茎干质量、根茎鲜质量比和根茎干质量比等指标影响显著,火箭盆容器和无纺布容器培育的苗木侧根系发达,根系质量好;而大田播种苗和普通塑料容器苗的主根发达,侧根少,根系质量较差。     

Influence of ground flora on Fraxinus mandshurica seedling growth on abandoned land and beneath forest canopy

A better understanding of root/shoot interactions influencing seedling growth on abandoned land could yield insight into seedling regeneration and restoration of the abandoned lands. Field work had been conducted for 2 years (2008–2009) to investigate the impacts of neighbouring plants on Manchurian Ash (Fraxinus mandshurica) seedling growth under the canopy of an old secondary forest and on the abandoned land exposed to full solar radiation in North-eastern China. Four different interaction treatments were designed for the study: neither shoot nor root interaction, shoot interaction only, root interaction only, and both shoot and root interaction. The presence of either the shoots or roots of neighbours had a competitive effect, reducing the growth of the target seedlings at each site. The total competitive effect of roots and shoots of neighbouring plants was significantly less than the sum of root and shoot competition separately on the abandoned land, but this difference was not significant beneath the forest canopy. Root competition was more restraining than shoot competition on the abandoned land. Target seedlings adjusted their root morphology and growth rates in response to the competitive effects from different parts of the neighbouring plants. Our results indicated that the root:shoot ratios of the target seedlings at the end of the experiment were affected by neither initial tree size nor the competition from either above- or belowground at either site. The results also highlight the importance of reducing root competition in boosting seedling regeneration and forest rehabilitation on the abandoned land.     

Direct seeding of late-successional trees to restore tropical montane forest

Natural regeneration of large-seeded, late-successional trees in fragmented tropical landscapes can be strongly limited by a lack of seed dispersal resulting in the need for more intensive restoration approaches, such as enrichment planting, to include these species in future forests. Direct seeding may be an alternative low-cost approach to planting nursery-raised tree seedlings, but there is minimal information on its efficacy or when in the successional process this technique will be most successful. We tested directly seeding five native tree species into habitats representing passive and active restoration approaches: (1) recently abandoned pasture; (2) naturally establishing, young secondary forests; and (3) young, mixed-species (fast-growing N-fixers and commercially valuable species) tree plantations established to facilitate montane forest recovery in southern Costa Rica. We monitored germination, survival, growth, and above- and below-ground biomass over a 2-year period. Germination in pastures, secondary forests, and tree plantations was similar (∼43%). Seedling survival after one and two years was significantly higher under tree plantations (91% year 1, 75% year 2) compared to secondary forests (76, 44%) or pastures (74, 41%). Moreover, seedlings had greater total biomass and lower root:shoot ratios in the plantations, suggesting higher nutrient availability in that treatment. Costs for direct seeding were 10- to 30-fold less per 100 seedlings after 2-year compared to nursery-raised seedlings planted at the same sites; however, there are important trade-offs to the two restoration approaches. Planting nursery-raised seedling is a more effective but higher cost approach for rapidly establishing canopy cover and restoring large areas whereas direct seeding is a more efficient way to enrich an existing system. We particularly recommend using direct seeding as a complimentary measure to the more intensive restoration approach of planting fast-growing and N-fixing trees.     

Regulating the Quality of Seedlings for Forest Restoration: Lessons from the National Greening Program in the Philippines

The use of low quality planting material is one of the major reasons for the limited success of past reforestation programs in the Philippines and elsewhere in the tropics. In the Philippines, a national policy has been in place since 2010, which regulates the quality of seedlings. As part of the policy, government reforestation programs are required to use only high quality seedlings from accredited seedling suppliers. A survey of nurseries producing seedlings for the National Greening Program in Eastern Visayas and Northern Mindanao regions was carried out to determine the effectiveness and challenges in implementing the forest nursery accreditation policy. The survey identified factors that limit the effectiveness of seedling quality regulation including lack of auditing of seedling quality in accredited nurseries, insufficient monitoring of the seedling supply chain among the network of nurseries supplying seedlings for reforestation programs, inadequate seedling production schedules, and inappropriate criteria for seedling quality assessment. The limited sources of high quality germplasm, nursery operators’ limited information on the attributes of high quality planting materials and lack of knowledge about high quality seedling production technologies contributed to the widespread production of low quality seedlings. The lack of seedling quality checks makes the government’s bidding scheme of seedling purchases prone to favouring the proliferation of low quality seedlings that are usually sold at lower prices. Nursery accreditation represents a major initiative in promoting the success of Philippine reforestation but our study found that considerable improvement of the policy and of its implementation is necessary. From our study, key lessons can be learned for the implementation of forest landscape restoration initiatives in other tropical developing countries.     

单叶省藤组培苗造林初步研究

单叶省藤(CalamussimplicifoliusC F Wei)是我国华南地区的优质商品藤种,其藤茎质量好,为编制家具的优质原料[1～3]。70年代以来的栽培试验表明,它不仅适应性强,而且速生,可广泛栽培于海南、广东、广西、云南和福建等省区,为我国棕榈藤栽培的首选藤种[1]。但野生资源由于长期过度采伐而枯竭,少量试验林未结实或结实甚少,推广栽培因种质资源奇缺而受到严重制约[2]。通过组织培养等生物技术工厂化快繁单叶省藤,是解决扩大栽培种苗问题的最好途径。因而开展了单叶省藤组培和造林技术研究,现将组培苗造林试验结果及其特殊性予以报道。1　…     

马尾松不同苗木类型与整地方式造林早期效果的研究

通过对不同苗木类型和整地方式的马尾松幼林的造林成活率、生长状况、生物量和根系特征进行测定，结果表明：容器苗可明显提高造林成活率、当年抽梢高和生物量；整地方式对造林当年的马尾松幼林有一定影响，而在第３年则对幼林树高和抽梢高起着明显促进作用。     

Root characteristics and growth potential of container and bare-root seedlings of red oak (<Emphasis Type="Italic">Quercus rubra</Emphasis> L.) in Ontario,Canada

Root characteristics and field performance of container and bare-root seedlings of red oak (Quercus rubra L.) were compared during the first growing season after planting. Sixty seedlings of each stock type were planted on a clearfell and weed-free site near Restoule, Ontario. Twenty-four additional seedlings from each stock type were compared at the start of the study in terms of shoot and root parameters. Measurement of root and shoot parameters were repeated at three dates during the first growing season in the field. The root systems of container stock had a larger number of first order lateral long roots and were significantly more fibrous than bare-root stock. These differences were sustained throughout the first growing season. In terms of field performance, container seedlings had 100% survival and achieved significant increases in both biomass and shoot extension. Bare-root seedlings suffered 25% mortality, significant shoot dieback and more variable growth. The mean relative growth rate (RGR) of container seedlings increased throughout the study period to a maximum of 30 mg/g/day, whereas the mean RGR of bare-root stock remained close to or below zero. Overall, the container seedlings proved less prone to transplanting shock than the bare-root seedlings, most likely due to favourable root architecture and the pattern of root development. Further work may be warranted in container design, growing regimes and root architecture to fully realise the potential of container systems for the production of high quality red oak seedlings across a range of site conditions.     

Root and shoot allometry of bareroot and container Douglas-fir seedlings

Detailed root and shoot development of bareroot and container Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedling stocktypes were compared during the first growing season after outplanting. The study was installed in raised beds with ideal environmental conditions and at a field reforestation site. Survival at both sites was 98% and did not differ between stocktypes. Seedlings were excavated in spring (5, 8, 12, and 16 weeks after planting) and in fall (35, 40, and 45 weeks after planting). In spring, container seedlings had more numbers of new roots and greater new root and shoot biomass than bareroot seedlings at both sites. In fall, bareroot seedlings consistently averaged more new root growth (though nonsignificant) than container seedlings suggesting that stocktype differences may not continue long-term. Container seedlings had significantly greater water percent than bareroot seedlings at the field site (all sample dates) and the raised bed site (weeks 5, 8, and 40 only). Regardless of environmental conditions or season, seedlings at both sites maintained water percent between 60 and 70% of fresh weight. Seedlings grown in the raised beds had much greater growth than those grown in the field. However, relative growth patterns for the two stocktypes were very similar on each site. The data generated establish baseline differences between stocktypes for root initiation, growth, and allometry during the first year after planting. Challenges associated with root development research are discussed.     

Physiological and field growth responses of oak seedlings to warm storage

The effect of warm storage (15°C) for 0, 15 or 31 days, applied after cold storage until April, and date of lifting to cold storage on the physiological condition and field performance of two-year-old oak seedlings (Quercus robur L.) was investigated. Assessments before planting included plant moisture status, root and shoot dry weight, root growth potential (RGP), while after planting root growth, shoot growth phenology, shoot and root dry matter accumulation and stem quality were assessed. Warm storage effects were large, but lift date effects were small. Warm storage for 31 days reduced height and diameter growth, stem quality, total biomass, root growth, and reduced stem quality in the field, but 15 days storage had a smaller effect. Warm storage delayed bud break and shoot growth cessation but survival was unaffected. The depletion of food reserves during storage and low moisture availability might have caused shoot dieback leading to the development of poor quality stems. There was evidence that dry weight fraction of both the shoot tip and the taproot provided good information on the quality of the stock before planting. RGP was also a good indicator of quality. Electrolyte leakage readings from fine and taproots were not reliable indicators of plant quality.     

Nutrient loading of forest tree seedlings to promote stress resistance and field performance: a Mediterranean perspective

The planting environment of Mediterranean areas is highly challenging as summer drought and winter frost jeopardize survival, and soil infertility limits establishment success. We review the potential for seedling nutrient loading to alleviate these post-planting stresses. A growing body of evidence indicates that nitrogen (N) rich seedlings have improved field performance in Mediterranean environments, due to their ability to grow new roots rapidly and out-compete weeds. In addition, frost resistance during hardening is crucial for relatively cold inland nurseries; recent research shows a positive relationship between N and shoot frost resistance though a knowledge gap exists regarding the influence of nutrition on root frost resistance. Some new evidence also implicates phosphorus as an important driver of seedling response in the Mediterranean due to its influence on root growth and physiology. Nutrient status influences other functional attributes critical to survival in Mediterranean areas, such as drought tolerance, root hydraulic conductivity, and mycorrhization. In light of the apparent benefits of high nutrient reserves for seedling performance in Mediterranean areas, we also review techniques for nursery nutrient loading. Exponential fertilization can be applied when species’ growth patterns match this application regime. However, many Mediterranean species exhibit episodic growth indicating that constant or fall fertilization could be more effective in reaching loading. In particular, late-season fertilization has shown good potential to avert nutrient dilution in the fall and increase frost resistance. Several needs for future research are identified, with special emphasis on the necessity to match fertilization regimes to species ecological traits and planting conditions.     

Effect of stock type characteristics and time of planting on field performance of aspen (Populus tremuloides Michx.) seedlings on boreal reclamation sites

Aspen (Populus tremuloides Michx) has great potential as a reclamation species for mining sites in the boreal forest, but planting stock has shown poor field performance after outplanting. In this study we tested how different aspen seedling characteristics and planting times affect field outplanting performance on reclamation sites. We produced three different types of aspen planting stock, which varied significantly in seedling size, root-to-shoot ratio (RSR), and total non-structural carbohydrate (TNC) reserves in roots, by artificially manipulating shoot growth during seedling production. All three stock types were then field-planted either in late summer, late fall, or early spring after frozen storage. Seedlings were outplanted onto two reclaimed open-pit mining areas in the boreal forest region of central and east-central Alberta, Canada, which varied significantly in latitude, reclamation history, and soil conditions. Overall, height growth was better in aspen stock types with high RSR and TNC reserves. Differences in field performance among aspen stock types appeared to be more strongly expressed when seedlings were exposed to more stressful environmental site conditions, such as low soil nutrients and moisture. Generally, aspen seedlings planted with leaves in the summer showed the poorest performance, and summer- or fall-planted seedlings with no shoot growth manipulation had much greater stem dieback after the first winter. This indicates that the dormancy and hardening of the stem, as a result of premature bud set treatments, could improve the outplanting performance of aspen seedlings, particularly those planted during summer and fall.     

Assessing the effect of late-season fertilization on Holm oak plant quality: insights from morpho–nutritional characterizations and water relations parameters

Forest restoration projects with Holm oak (Quercus ilex) have had limited success, mostly due to water stress after planting and poor plant quality. Recent studies indicated that large and nutrient rich plants perform better in Mediterranean areas, suggesting that late-season fertilization may improve plant quality and field performance. The purpose of this study was to assess the effect of late-season fertilization on the quality of Holm oak seedlings, as determined by morphological, nutritional, and water relations analyses. We grew Holm oak nursery seedlings under 5 different late-season fertilization regimes and then analyzed morphological characteristics, nutritional status, and water relations parameters of the fertilization groups at the end of the nursery period. We also analyzed the effect of fertilization on nutritional status by use of vector nomograms. Our results indicated that late-season NPK fertilization improved shoot and root growth, and the overall nutritional status of seedlings. The lack of late-season fertilization leads to nutrient deficiency in plants, whilst the application of imbalanced fertilization treatments may trigger nutrient luxury consumption and nutrient dilution, pointing out the importance of NPK proportions in the fertilizer. Moreover, late-season fertilization with nitrogen might improve the drought resistance of seedling by enhancing their osmotic adjustment.     

Regeneration of Picea abies forests on highly productive peatlands—clearcutting or selective cutting?

The study was focused on severe forest regeneration problems that are typical for highly productive peatlands. The aim of the study was to give recommendations for practical forestry on how to renew the forests. Experiments with different forest regeneration methods on highly productive peatlands were set up in nine mature Norway spruce (Picea abies (L.) Karst.) forests in northern, central and southern Sweden. The treatments in the study were natural regeneration in shelterwoods (at densities of 140 and 200 stems ha^‐1), planting of bare‐root spruce seedlings after site preparation (mounding) in the shelterwoods and on clearcuts, and planting without site preparation on clearcuts.

Judging from the extent of windthrow in the denser shelterwoods and the stocking of natural regeneration under remaining shelter trees 4–5 years after final cut, shelterwood regeneration is a promising method. On clearcuts, planting without site preparation resulted in poor seedling survival, large extent of damage to the seedlings and small height increment. Planting in mounds on the clearcuts was much more successful. The best survival of planted seedlings was observed in the shelterwoods. Both the denser and the sparse shelterwoods seemed to give satisfactory protection to the seedlings.

Final recommendations on the most suitable forest renewal method(s) will be made in about five years after the removal of the shelter trees.     

草珊瑚不同播种育苗方式效果研究

试验采用营养袋播种育苗和田间苗床播种育苗2种方式进行草珊瑚苗木生长量的比较研究,结果表明：2种育苗方式对苗木的出土时间没有影响,但对种子的发芽率具有极显著影响,前者比后者提高29.3%。2种不同育苗方式的高径生长量、根系生长量及总生物量（鲜重）差异极显著;营养袋有利于苗木的高径生长和主根生长,总生物量比田间苗床播种苗提高30.4%;但侧根生长相比较差。为提高移栽成活率,在生产上提倡营养袋育苗。     

Seedling quality and field performance of commercial stocklots of containerized holm oak (Quercus ilex) in Mediterranean Spain: an approach for establishing a quality standard

Holm oak is the hardwood most used for reforestation in Mediterranean Spain, which makes the development of stock quality standards in order to improve establishment success, a priority. However, its nursery culture is characterized by a wide range of practices resulting in stock heterogeneity and a potentially varied outplanting performance. Previous research has focused on specific seedling quality attributes, obviating the integral effect of nursery culture on overall quality. We studied growing regime, seedling quality, and field performance in nine holm oak stocklots produced in commercial nurseries during two consecutive growing years. Results proved variations in field performance were related to stocklot quality and, hence, to the growing regime practised. This dependence on stock quality may vary with planting site weather: in the drier year, survival was related to attributes like height, water status and K concentration, while, in the second, milder year, only growth performance was related to nutrient concentrations, plant size and water status. Results indicated the following quality standards for height: 12–17 cm, diameter: 3.5–4.8 mm, shoot and root weights: 1.3–1.6 and 2.8–4.7 g, respectively, N–P–K foliar concentrations: over 10–0.9–3.7 mg g⁻¹, respectively and in water status parameters: E_MX < 5 MPa and SWD_TL > 15%. These attributes can be adjusted using nursery cultural practices in order to meet seedling quality standards for holm oak for planting across similar sites.     

Assessing the Quality of Seedlings in Small-scale Nurseries in the Highlands of Cameroon: The Use of Growth Characteristics and Quality Thresholds as Indicators

In developing countries seedlings are often produced in small-scale nurseries as a means to raise tree planting materials and, provided they are of high quality, these can form the basis for successful forest plantation initiatives. This study uses morphological characteristics to assess the quality of seedlings of three tree species (Persea americana, Cola acuminata and Dacryodes edulis) in the Western Highlands of Cameroon. The growth characteristics and quality indicators of seedlings produced in two nursery categories were compared—three nurseries collaborating with the tree domestication program implemented by the World Agroforestry Centre (Category 1) and three nurseries collaborating with government and non-governmental projects (Category 2). Seedlings produced in nurseries in the first nursery category were found to be uniform in terms of growth characteristics and quality indicators. In the second category, there were relatively higher seedling proportions having measured parameters out of threshold standards for root to shoot ratio and for sturdiness quotient. Significant differences between nursery categories (at α = 0.05) were detected in root collar diameter of C. acuminata and D. edulis and in shoot dry weight of D. edulis. These differences could be attributed to better nursery management as a results of training and technical backstopping received by nursery operators in Category 1. The level of experience of nursery operators in both categories was found to be insufficient and greater exposure to skills and knowledge could enhance the production of high quality seedlings.     

Ecophysiological variation in two provenances of Pinus flexilis seedlings across an elevation gradient from forest to alpine

Climate change is predicted to cause upward shifts in forest tree distributions, which will require seedling recruitment beyond current forest boundaries. However, predicting the likelihood of successful plant establishment beyond current species' ranges under changing climate is complicated by the interaction of genetic and environmental controls on seedling establishment. To determine how genetics and climate may interact to affect seedling establishment, we transplanted recently germinated seedlings from high- and low-elevation provenances (HI and LO, respectively) of Pinus flexilis in common gardens arrayed along an elevation and canopy gradient from subalpine forest into the alpine zone and examined differences in physiology and morphology between provenances and among sites. Plant dry mass, projected leaf area and shoot:root ratios were 12-40% greater in LO compared with HI seedlings at each elevation. There were no significant changes in these variables among sites except for decreased dry mass of LO seedlings in the alpine site. Photosynthesis, carbon balance (photosynthesis/respiration) and conductance increased >2× with elevation for both provenances, and were 35-77% greater in LO seedlings compared with HI seedlings. There were no differences in dark-adapted chlorophyll fluorescence (Fv/Fm) among sites or between provenances. Our results suggest that for P. flexilis seedlings, provenances selected for above-ground growth may outperform those selected for stress resistance in the absence of harsh climatic conditions, even well above the species' range limits in the alpine zone. This indicates that forest genetics may be important to understanding and managing species' range adjustments due to climate change.