Is light the key factor for success of tube shelters in forest restoration plantings under Mediterranean climates?

Tube shelters were designed to protect against browsing, but they improve seedling survival in Mediterranean dry climates. Mechanisms for this response, however, are not fully understood and this knowledge can be useful to help design optimal tube shelters for Mediterranean species and climates. Our objective in this study was to determine if the positive effect of tube shelters is due to enhanced growth during the wet season or to reduced light stress during the dry season. We performed two independent experiments. In the first, we assessed root growth during the wet season in two Mediterranean species with contrasting light tolerance (Quercus ilex L. and Pinus halepensis Mill.) growing in tubes with varying light transmissivity. In the second experiment, we studied the response of a Quercus ilex plantation to different shelter treatments. Root growth during the wet season was reduced with decreasing light transmissivity in the shade intolerant P. halepensis, but not in the shade tolerant Quercus ilex. Survival of Q. ilex shaded by a mesh shelter only during summer was higher than in unsheltered seedlings and similar to the survival in tube and mesh shelters during the whole season. This suggests that shade during the dry period was the main factor explaining survival in this species. This effect could be related to the lower leaf temperature recorded in sheltered seedlings. We conclude that Q. ilex (and perhaps other late successional, shade tolerant Mediterranean species) should be planted in tubes with the currently used light transmissivity because these shelters reduce light stress in summer without impairing root growth in the wet season. However, current tubes impair root growth in P. halepensis (and likely other pioneer, shade intolerant Mediterranean species), so higher transmissivity tubes may be necessary. Optimal transmissivity for tube shelter in Mediterranean climates is species-specific and identifiable as the point that minimizes light stress during summer without impairing root growth in the wet season.     

Effects of nursery shading on seedling quality and post-planting performance in two Mediterranean species with contrasting shade tolerance

In Mediterranean climates, seedlings are frequently shaded in the nursery to avoid heat damage and save water. However, the impact of this shading on the seedling quality and transplanting performance of Mediterranean species is not well known. We studied the effect of nursery shading on pre-planting features and post-planting performance of two Mediterranean tree species: the shade-intolerant pioneer Pinus halepensis and the shade-tolerant late-successional Quercus ilex. We grew one-year-old seedlings of both species under 100, 40 and 5% full sunlight. Shade had a low impact on the morphology and physiology of Q. ilex seedlings. In pines, only the deep shade treatment produced low quality seedlings with poor root development. In both species, transference to high light at planting in autumn did not impose any additional stress than that caused by frosts, but initial root growth was impaired in the two shaded treatments in pine. Post-planting growth and survival of oak seedlings showed no difference between treatments. Pine seedlings grown in deep shade showed higher mortality and lower growth after planting than those grown in full sun and intermediate light treatments, while intermediate light only reduced growth. For the nursery culture of Q. ilex seedlings, we advise using low light levels during summer to save water without impairing field performance. In P. halepensis, seedlings should be cultured under full sunlight conditions to maximize post-planting growth, but they can be cultured under intermediate light without impairing survival.     

Refugial forests of the southern Appalachians: photosynthesis and survival in current-year Abies fraseri seedlings

Fraser fir (Abies fraseri (Pursh) Poiret) is an endemic, high-elevation conifer confined to six relict mountaintop communities in the southern Appalachian Mountains, USA. High adult mortality has occurred over the past 50 years, possibly the result of an introduced insect (Adelges piceae Ratzeburg), air pollution, or both. Knowledge of the mechanisms of and limitations to seedling establishment may allow reestablisment and perpetuation of this unique community type, notwithstanding global climate change. We monitored seedling emergence and mortality in relation to photosynthetic performance and water relations in microsites differing in canopy openness (sunlight exposure) over the summer of 2004. Abundance of cotyledonous seedlings in early summer was 2.3 times greater (849 versus 366 seedlings m(-2)) in microsites with lower sky exposure (greater canopy closure) than in microsites with greater sky exposure (greater canopy openness). In contrast, late-season abundance and survival were greater in areas beneath more open canopies than in areas beneath less open canopies (3.3 times and 11.7 times greater, respectively). However, newly emerged seedling survival in a completely open site (no overhead canopy) was zero, despite an initial density of 124 seedlings m(-2). Seedling water status was similar in open- and closed-canopy sites (-0.52 and -0.74 MPa, respectively). Photosynthetic carbon gain was higher in newly emerged seedlings at open canopy than at closed canopy sites, especially during early morning. Based on photosynthetic light response curves and measured sunlight regimes, seedlings in open canopy sites were estimated to assimilate 3.3-4.5 times more carbon than seedlings at closed sites. Reductions in carbon gain of closed-site seedlings, primarily a result of limited sunlight, corresponded to substantial increases in seedling mortality (98 versus 79% in open canopy sites). Thus, sunlight exposure, which reflects overstory canopy structure, appears to be an important factor influencing newly emerged seedling survival and distribution.     

Shade tolerance, photoinhibition sensitivity and phenotypic plasticity of Ilex aquifolium in continental Mediterranean sites

Shade tolerance, plastic phenotypic response to light and sensitivity to photoinhibition were studied in holly (Ilex aquifolium L.) seedlings transported from the field to a greenhouse and in adult trees in the field. All plants were growing in, or originated from, continental Mediterranean sites in central Spain. Seedlings tolerated moderate but not deep shade. Mortality was high and growth reduced in 1% sunlight. Survival was maximal in 12% sunlight and minimal in full sunlight, although the relative growth rate of the seedlings surviving in high light was similar to that of plants in moderate shade. Maximum photochemical efficiency at predawn was significantly lower in sun plants than in shade plants in the field, revealing chronic photoinhibition that was most pronounced in winter. Plasticity in response to available light varied according to the variable studied, being low for photosynthetic capacity and stomatal conductance, and high for specific leaf area, root:shoot ratio and leaf area ratio, particularly in seedlings. Differences in water relations and hydraulic features between sun and shade plants in the field were marginal. High water potential at the turgor loss point of field-grown plants suggested that holly is sensitive to drought during both the seedling and the adult stage. Low relative growth rates in both high and low light with low physiological plasticity in response to light indicate the existence of a stress-tolerance mechanism. We conclude that holly is a facultative understory plant in areas of oceanic and relatively mild climate, but an obligate understory plant in dry continental areas such as the study site. The impact of abandonment of traditional management practices and climate change on these Mediterranean populations is discussed.     

Effects of light availability and soil flooding on growth and photosynthetic characteristics of Genipa americana L. seedlings

Genipa americana L. (Rubiaceae) is a late successional neotropical fruit tree used in riparian forest restoration programs. We analyze the effects of light availability and soil flooding on the growth and photosynthetic characteristics of G. americana seedlings under nursery and natural light conditions. Two light levels (full sunlight and shade), and two levels of soil water (flooded and control) were used in the experiment. Flooding induced significant changes in the total seedling biomass (P < 0.01). The differences among water treatments were 70 and 10% at full sunlight and shade, respectively. These changes were explained by alterations in the maximum quantum efficiency of the photosystem 2 (Fv/Fm), light-saturated net photosynthetic rates (A _sat) and intrinsic water use efficiency (WUE). G. americana has high physiological plasticity in relation to the light availability and flooding, although significant interactive effects between high light exposure and soil flooding were observed in several photosynthetic and growth variables. The results highlight the importance of the synergistic effects between physical environmental variables on the establishment and growth of G. americana seedlings. In the practical point of view we can indicate that, in degraded riparian forests subjected to soil flooding, seedlings of this species should be planted under partially shaded environments.     

遮阳网遮光度对红椎大田播种育苗效果的影响

设置了4种不同的遮光处理:全光照、遮光度40%、遮光度60%、遮光度70%,探讨不同遮光度对红椎大田播种育苗苗木保存率和苗木生长的影响。结果表明:遮光度对红椎大田播种育苗苗木保存率和苗木生长有显著影响。遮光度达70%时,其苗木保存率、苗木高生长与地径生长均表现最佳。     

Differential light responses of Mediterranean tree saplings: linking ecophysiology with regeneration niche in four co-occurring species

The ecophysiological mechanisms underlying plant-plant interactions and forest regeneration processes in Mediterranean ecosystems are poorly understood, and the experimental evidence for the role of light availability in these processes is particularly scant. We analyzed the effects of high and low irradiances on 31 ecological, morphological and physiological variables in saplings of four late-successional Mediterranean trees, two deciduous (Acer opalus subsp. granatense (Boiss.) Font Quer & Rothm. and Quercus pyrenaica Willd.) and two evergreen (Pinus nigra Arnold subsp. salzmannii (Dunal) Franco and Quercus ilex L.), which coexist in mature montane forests. Species differed in both their capacity to withstand high radiation and in their shade tolerance. The two deciduous species were the least tolerant to high radiation, exhibiting both dynamic and chronic photoinhibition in full sunlight, with severe implications for gas exchange and photosynthetic performance. Excess light severely limited the survival of A. opalus subsp. granatense, even minor reductions of excessive radiation (from full sunlight to 80% sunlight) being crucial for sapling survival. Among species, P. nigra was the most tolerant of high irradiances but showed the poorest shade acclimation. Interspecific differences in the mechanisms of response to light provide a partial explanation of the differential regeneration patterns previously reported for these species, with shade-tolerant plants (i.e., deciduous broadleaf species) benefiting the most from associations with nurse plants. We conclude that light availability is an important environmental factor defining the regeneration niche of Mediterranean woody species.     

Testing Biodegradable Seedling Containers as an Alternative for Polythene Tubes in Tropical Small-Scale Tree Nurseries

Polythene tubes are the most commonly used seedling containers and their adoption can be attributed to high water retention that enhances seedling establishment as well as the desire for low-cost readily-available containers by nursery operators. Polythene tubes have drawbacks, however, because they adversely affect seedling root growth and are an environmental hazard. This study was conducted in Meru, Eastern Kenya, to investigate whether small-scale tree nursery operators are likely to adopt biodegradable seedling containers (cellulose papers and banana sheaths). It was hypothesised that biodegradable containers are better for seedling growth and are more environmental friendly than the widely used polythene bags. The study assessed the frequency of watering and growth (height and basal diameter) of Calliandra calothyrsus seedlings produced in various biodegradable containers under three conditions with varying watering requirements, i.e. light tree shade, shade net and polythene chambers, the first being widely used by farmers. The performance of these seedlings was later monitored in the field. Seedlings produced in biodegradable containers required more frequent watering than those in polythene bags under light tree shade and shade nets but less frequent in polythene chambers. Seedlings produced in polythene tubes had higher growth rates in the nursery, but when transplanted to the field, they were overtaken by those grown in the biodegradable containers due to transplanting shock after the polythene containers were removed. Biodegradable seedling containers can therefore be adopted in areas where water is not very limiting, and evaporation rates could be reduced and water-use efficiency improved by raising seedlings in simple polythene structures.     

Establishment and early growth of introduced indigenous tree species in typical plantations and shrubland in South China

Unsuccessful colonization by indigenous tree species into established plantations has retarded the succession of artificial plantations to more natural, secondary forests in South China. To understand how to improve colonization by seedlings of indigenous species, we determined how performance of indigenous seedlings is affected by seedling species (the shade-intolerant Castanopsis chinensis, the moderately shade-intolerant Michelia chapensis, and the shade-tolerant Psychotria rubra), the site into which the seedlings were transplanted (a mixed-legume plantation, a eucalyptus plantation, a mixed-native plantation, a mixed-conifer plantation, and a shrubland), and site preparation (removal or retention of understory vegetation and litter). Seedling survival and growth were generally increased by removal of understory vegetation and litter. C. chinensis and M. chapensis grew better in the mixed-legume and mixed-conifer plantations, while P. rubra grew better in mixed-native and mixed-conifer plantations. Responses of the transplanted seedlings to environmental factors were species specific. The effects of light on seedling survival and growth were correlated with the shade tolerance of the species. Soil moisture was important; it was positively correlated with survival but negatively correlated with growth of C. chinensis seedlings. Growth of C. chinensis and M. chapensis was positively correlated with soil potassium, while growth of P. rubra was positively correlated with soil organic matter but negatively correlated with soil hydrolyzed nitrogen. These findings suggest that we should select suitable native species under the different plantations before improvement of plantations. Light and soil moisture are most important environmental factors for the selection of species specific. Site preparation and fertilizer are needed during the improvement of those plantations.     

伯乐树1年生幼苗的光响应特征

采用自然全光照的16.72%(s17)、24.51%(s25)、36.49%(s36)和100%(s100)及林地下14.02%(s14)等5种光照强度,比较分析不同光照条件下的环境因子日变化、1年生幼苗成活率、生长量等指标,以期揭示幼苗对于夏季不同光照条件的光响应特征。结果表明:不同光照强度下,环境因子日变化显著不同,幼苗的成活率、形态和生理指标等均存在显著差异。极度弱光下(s17)幼苗生长迟缓;而全光照环境(s100)和较强光照下(s36)易造成幼苗的死亡;仅适度遮荫(s25)适于幼苗的生长;林下栽培(s14)的相对光照强度虽然较弱,但其各种小气候因子均明显优于无林地的其它光照条件,幼苗成活率、水分利用和平均净光合速率等指标均达到最优。建议抚育幼苗时应采用适宜的遮荫处理,可选择在中等郁闭度的林下(自然遮荫)或在大田中采用夏季单层遮荫。     

Conspecific negative density dependence in a long-lived conifer,yew <Emphasis Type="Italic">Taxus baccata</Emphasis> L.

Conspecific negative density dependence in tree species can have major consequences for structuring of communities, yet in temperate forests this phenomenon remains largely unexplored. We investigated density-dependent recruitment in the long-lived conifer yew (Taxus baccata L.), for which low levels of regeneration have been linked with reduced light availability beneath conspecific canopies and speculative autotoxic effects. We combined in situ assessment of yew regeneration in a temperate forest in southwest Ireland with ex situ pot experiments to assess whether light and/or beneath conspecific canopy conditions inhibit the germination and early growth of yew. In field experiments, recruitment was related to distance from adult conspecifics, with higher levels of regeneration found in areas not beneath yew canopies. However, reduced light availability beneath the dense yew canopy was not responsible for this pattern, with abundant seedlings found in areas of equivalent or even lower light availability. Pot experiments showed that seedling survival was highest in deep shade treatments. Experiments also demonstrated that the addition of yew needles negatively affected seedling growth, but not seed germination. Together, our results suggest that under natural forest conditions, the absence of regeneration beneath mature yew canopies, often attributed to low light availability, may at least be partly related to substances in yew canopy or leaf leachates.     

Growth and photosynthetic responses of four Virginia Piedmont tree species to shade

To determine the effects of shade on biomass, carbon allocation patterns and photosynthetic response, seedlings of loblolly pine (Pinus taeda L.), white pine (Pinus strobus L.), red maple (Acer rubrum L.), and yellow-poplar (Liriodendron tulipifera L.) were grown without shade or in shade treatments providing a 79 or 89% reduction of full sunlight for two growing seasons. The shade treatments resulted in less total biomass for all species, with loblolly pine showing the greatest shade-induced growth reduction. Yellow-poplar was the only species to show increased stem height growth in the 89% shade treatment. The shade treatments increased specific leaf area of all species. Quantum efficiency, dark respiration and light compensation point were generally not affected by the shade treatments. Quantum efficiency, dark respiration, maximum photosynthesis and light compensation point did not change consistently between the first and second growing seasons. We conclude that differences in shade tolerance among these species are not the result of changes in the photosynthetic mechanism in response to shade.     

Interactive responses of Quercus suber L. seedlings to light and mild water stress: effects on morphology and gas exchange traits

? The combined effect of water stress and light on seedlings of forest species is a key factor to determine the best silvicultural and afforestation practices in the Mediterranean area.

? The aims of this work was (1) to determine the optimal light level for the early development of cork oak seedlings under mild water stress and (2) to test if the combined effect of water stress and light followed the trade-off, the facilitation or the orthogonal hypothesis.

? Shade reduced instantaneous photosynthetic rates and water use efficiency in cork oak. However, seedlings grown under moderate shade (15% of full sunlight) were capable to accumulate similar amount of biomass than those grown under more illuminated environments by increasing their specific leaf area. Absolute differences in net photosynthesis between light treatments were higher in well watered than in water stressed seedlings. However, the impact of both factors on overall growth was orthogonal.

? We concluded that cork oak development is impaired under deep shade (5% of full sunlight) but it can be optimal under moderate shade (15% of full sunlight) even under moderate water stress. Implications of these patterns on regeneration, cultivation and afforestation of cork oak are discussed.

     

遮荫处理对臭椿幼苗早期生长的影响

在不同光照梯度(55.44%自然全光照、21.12%自然全光照、3.47%自然全光照和0.86%自然全光照)的人工遮荫条件下,研究了不同光照强度对臭椿Ailanthus altissima幼苗早期生长的影响。结果表明:移栽后约3个月的臭椿幼苗株高、基径、复叶数、主根长、单株叶面积、总干质量、根冠比、相对生长率和净同化率等生长参数均以55.44%自然全光照处理最大,分别为77.30 cm、13.82 mm、17.80、67.20 cm、4 852.22 cm2、60.57 g、0.24、49.76×10-3g.d-1和5.62×10-5g.cm-2d-1,而且随着光照强度的减弱而减小。研究结果可为臭椿苗木繁育与造林实践提供参考。     

红皮云杉幼树不同光环境下生理生态特征研究

为了探讨红皮云杉光舍特征与树木枝叶分布格局，生物量及其分配的关系，采用人工控制光照环境，在4个光照条件下进行红皮云杉生长试验，结果表明，红皮云杉幼树瞬时最大净光合速率以透光70％处理最大，10％处理最小，对数据进行方差分析，结果差异显著；并且净光合速率最大的处理其暗呼吸速率也最大，光补偿点和光饱和点也随透光率而变化，透光率小(10％)的处理光补偿点也小，说明云杉幼树对庇荫环境的适应；不同光照条件下幼树的形态特征也表现出明显差异，长期生活在相对较暗的光环境中，形成细而高的树干，侧枝数量增加、长度加长，侧枝角度增大；云杉小苗(9年生以前)以相对照度70％处理生物量最大，9年生以后以全光区生物量最大，说明云杉在9年生以前耐庇荫，9年生以后喜光。     

遮光对野鸦椿幼苗生长和光合生理特性的影响

为了解不同光照条件下野鸦椿幼苗的生长发育状况和光合生理特性,探讨野鸦椿幼苗生长发育的最佳光照条件.采用遮光方式模拟不同光照强度(全光照,遮光30％、60％和90％),研究野鸦椿幼苗的生长、生物量、光合色素含量、生理指标和光合日变化等光合特性的响应规律.结果表明:随着遮光程度的增加,野鸦椿幼苗的生长量均高于对照,以遮光6...     

Shading System on Sandalwood Seedlings in Timor, East Nusa Tenggara, Indonesia

Mortality rates of seedlings of sandalwood (Santalum album L.) of more than 40% have been observed in cool wet upland areas in various countries. Mortalities are caused by low sunlight intensity and high soil humidity after watering, when seedlings are growing in modern shade houses (paranet roof with 50% solar intensity) in relatively high rainfall upland areas. To cope with this problem, it is proposed to use conventional sheltering construction with roof material modification as one of the alternatives to implemented to intensity solar radiation and reduce soil humidity. This method was proposed to increase the growth rate and reduce fungal decay in seedlings. The study observed the effect of various roof materials used with conventional nursery construction on growth of sandalwood seedlings. A randomized block design was adopted, with treatments consisting of roof material types, including coconut leaf (Cocus nucifera), imperata grass (Imperata cylindrica), transparent plastic, plastic roof (laserlaip), and paranet. A modern shade house was used as a control variable. The experimental design included three blocks and each block consisted of 50 replications. Conventional sheltering devices were found to be superior to other devices for 8-month-old sandalwood seedling. The conventional seedling sheltering device from corrugated plastic roof (laserlaip) resulted the greatest growth in height and diameter and greatest survival percentage of sandalwood seedlings. The highest to lowest rank of growth in order were plastic roof, paranet, transparent plastic, coconut leaf, imperata grass, and shade house, with survival percentage in each sheltering device of 62.12%, 53.55%, 41.81%, 40.82%, 38.51%, and 12.82%, respectively. Waving plastic roof resulted in highest seedling growth than other treatment because it is protected from rain water (waterlogging) and will raise the temperature and the intensity of sunlight received.     

Transplantation of Larix principis-rupprechtii Mayr. and Picea meyeri Rehd. seedlings to low altitude and two contrasting light environments reveals climate warming effects on early seedling performance

By using altitude as a surrogate of warming treatment, we transplanted seedlings of Larix principis-rupprechtii Mayr. and Picea meyeri Rehd. Et Wils. from their original altitude (2000?m a.s.l.) to a lower altitude (1500?m a.s.l.) and grown in two light environments (full and low) at both altitudes for two years in north-central China, to determine the effects of warming and light on seedling survival and growth performance. The results showed that altitude and light did not affect survival rates of both species, but had significant interactions on their growth performance. Although both species had greater heights, root collar diameters and biomass accumulations in full than in low light at high altitude, such growth advantage disappeared or even reversed at low altitude, especially for L. principis-rupprechtii. Moreover, L. principis-rupprechtii seedlings increased specific leaf area, leaf area ratio and shoot:root ratio at low altitude. Despite this study is relatively short termed, such results demonstrated the importance of shade tolerance in species responses to warming and may imply an altered regeneration pattern of L. principis-rupprechtii if the current warming trend remains in the future.     

Acclimation abilities of three tropical rainforest seedlings to an increase in light intensity

Current practices in many nurseries involve the germination of tropical rainforest seedlings in shaded conditions and transferral, at a later date, to environments with greater light intensities. Determination of the ability of these seedlings to acclimate to increased light intensities will allow seedling stock to be processed with maximum efficiency within the nursery. The acclimation abilities of three species, Agyrodendron actinophyllum, Cardwellia sublimis, and Flindersia brayleyana, commonly found in north Queensland's rainforests were investigated in this study. These particular species are highly valued for their cabinet timber qualities and are being reared in nurseries for use in reforestation trials and programs in north Queensland. Seedlings were initially raised in greenhouse conditions under two layers of shade cloth (16% of full sunlight) and then transferred into full sunlight at three different ages (3 weeks, 10 weeks, 14 weeks). Upon transfer, organ ratios and the direction of dry matter distribution was determined for each species and age group. Approximately 3 months after the seedlings were transferred, the acclimation ability of each species and age group was then determined. Dry matter distribution was found to change with age, irrespective of light environment. Individuals within a species with larger root systems and thicker or more dense leaves had a greater acclimation ability than those with smaller root systems and thinner or less dense leaves. Furthermore, individuals within a species whose dry matter distribution upon transfer was directed towards developing a large root system, and a small photosynthetic area and mass, had a greater acclimation ability than those whose dry matter distribution was directed away from such morphologies. Awareness of these relationships allows a better understanding of seedling response to gap formation in natural forests, and also allows plant nursery operators to make a more informed decision about when to move seedlings to environments with a higher light intensity.     

Morphological and photosynthetic alterations in the Yellow-ipe, <Emphasis Type="Italic">Tabebuia chrysotricha</Emphasis> (Mart. Ex DC.) Standl., under nursery shading and gas exchange after being transferred to full sunlight

The objective of this study was to evaluate the effect of nursery shading on the Yellow-ipe seedling (Tabebuia chrysotricha) growth, photosynthesis, and photosynthetic acclimation after being transferred into direct sunlight. The Yellow-ipe seedlings were grown under 0, 50, 70 and 95% shade. At the 134th day of sowing, leaf gas exchange and chlorophyll were measured under current growth shading, after exposure to 15 min and two day full sunlight. With the increase of shading, the Yellow-ipe seedlings allocated more biomass to the stem and leaves and less to the roots, and there was an increase in the leaf area ratio and specific leaf area. In relation to 0% of shading there was a increase of 211% in stem, 116% in leaf, and a reduction of 200% in roots biomass when seedling were grown under 95% of shading. The total biomass accumulation had a high correlation with collar diameter (r = 0.96). More than 70% of the shading reduced the photosynthesis, and after transferring the seedlings into full sunlight, more than 50% of the shading induced a reduction in chlorophyll, stomatal conductance, photosynthesis and instantaneous carboxylation efficiency, suggesting the presence of a photoinhibition process. The Yellow-ipe seedling growth under nursery conditions should not be done under more than 50% shading, which may result in the lower seedling quality and poorer acclimation to transplantation, particularly to severe degraded areas with direct sunlight. The species can be used for recovering from totally devastated forest areas to initial recovery when full canopy are forming.