Environmental factors regulating the radial oxygen loss from roots of Myriophyllum spicatum and Potamogeton crispus

Myriophyllum spicatum and Potamogeton crispus are common species of shallow eutrophic lakes in north-eastern Germany, where a slow recovery of the submersed aquatic vegetation was observed. Thus, the characterisation of the root oxygen release (ROL) as well as its implication for geochemical processes in the sediment are of particular interest. A combination of microelectrode measurements, methylene blue agar and a titanium(III) redox buffer was used to investigate the influence of the oxygen content in the water column on ROL, diel ROL dynamics as well as the impact of sediment milieu. Oxygen gradients around the roots revealed a maximum oxygen diffusion zone of up to 250 μm. During a sequence with a light/dark cycle as well as alternating aeration of the water column, maximum ROL with up to 35% oxygen saturation at the root surface occurred under light/O₂-saturated conditions. A decrease to about 30% was observed under dark/O₂-saturated conditions, no ROL was detected at dark/O₂-depleted conditions and only a weak ROL with 5–10% oxygen saturation at the root surface was measured under light but O₂-depleted water column. These results indicate, that during darkness, ROL is supplied by oxygen from the water column and even during illumination and active photosynthesis production, ROL is modified by the oxygen content in the water column. Visualisation of ROL patterns revealed an enhanced ROL for plants which were grown in sulfidic littoral sediment in comparison to plants grown in pure quartz sand. For both plant species grown in sulfidic littoral sediment, a ROL rate of 3–4 μmol O₂ h⁻¹ plant⁻¹ was determined with the Ti(III) redox buffer. For plants grown in pure quartz sand, the ROL rate decreased to 1–2 μmol O₂ h⁻¹ plant⁻¹. Hence, aside from the oxygen content in the water column, the redox conditions and microbial oxygen demand in the sediment has to be considered as a further major determinant of ROL.     

Chemical profile of the North American native Myriophyllum sibiricum compared to the invasive M. spicatum

Myriophyllum spicatum L. is a nonindigenous invasive plant in North America that can displace the closely related native Myriophyllum sibiricum Komarov. We analyzed the chemical composition (including: C, N, P, polyphenols, lignin, nonpolar extractables, and sugars) of M. spicatum and M. sibiricum and determined how the chemistry of the two species varied by plant part with growing environment (lake versus tank), irradiance (full sun versus 50% shading), and season (July through September). M. spicatum had higher concentrations of carbon, polyphenols and lignin (C: 47%; polyphenols: 5.5%; lignin: 18%) than M. sibiricum (C: 42%; polyphenols: 3.7%; lignin: 9%) while M. sibiricum had a higher concentration of ash under all conditions (12% versus 8% for M. spicatum). Apical meristems of both species had the highest concentration of carbon, polyphenols, and tellimagrandin II, followed by leaves and stems. Tellimagrandin II was present in apical meristems of both M. spicatum (24.6 mg g⁻¹ dm) and M. sibiricum (11.1 mg g⁻¹ dm). Variation in irradiance from 490 (shade) to 940 (sun) μmol of photons m⁻² s⁻¹ had no effect on C, N, and polyphenol concentrations, suggesting that light levels above 490 μmol of photons m⁻² s⁻¹ do not alter chemical composition. The higher concentration of polyphenols and lignin in M. spicatum relative to M. sibiricum may provide advantages that facilitate invasion and displacement of native plants.     

Uptake of stable N isotopes by Myriophyllum spicatum is not selective

Preference for ¹⁴N over ¹⁵N was assessed for Myriophyllum spicatum L. by exposing shoots to the same initial concentration of N, but different isotopic compositions (100% ¹⁴N, 50% ¹⁴N and 50% ¹⁵N, and 100% ¹⁵N) for 6 weeks. Regardless of treatment, all available inorganic N was taken up by M. spicatum at similar rates. Significant differences in tissue isotopic composition were evident across treatments in 2 weeks, and persisted for the remainder of the experiment. At 6 weeks, atom (at) % ¹⁵N was twice as high in the 100% ¹⁵N treatment as in the 50% ¹⁵N treatment, and the at% of the ¹⁴N treatment (0% ¹⁵N) remained unchanged. This observed absence of preference for ¹⁴N in M. spicatum suggests that this species can be useful in determining N sources to freshwater systems, which often differ in ¹⁵N/¹⁴N ratio.     

太湖蓝藻死亡腐烂产物对狐尾藻和水质的影响

为揭示富营养化造成淡水生态系统中沉水植物消亡的机制,本文就太湖蓝藻大规模死亡腐烂后的产物对水体水质以及沉水植物造成的影响进行了研究。本文以沉水植物穗花狐尾藻为研究对象,测定了水质参数（pH值,浊度（NTU）,温度,溶解氧（DO）,电导率（Ec）,盐度,磷酸根（PO43-））以及狐尾藻Fv/Fm指标。将采自太湖梅梁湾的蓝藻水华,降解一个星期左右。随后在处理组中加入该蓝藻降解物,对照组中不加入。结果表明,加入蓝藻死亡腐解液后,水体的溶解氧、pH值均比对照组显著降低,浊度则显著升高。对照组中穗花狐尾藻生长良好,其Fv/Fm约为0.8,而经蓝藻腐烂液处理的实验组,穗花狐尾藻在两天之内便接近死亡状态,其Fv/Fm值降至0.3左右。以上结果表明蓝藻死亡腐烂后形成的腐解液确实能在很短时间内给沉水植被带来灭顶之灾。而在短时间内并不能形成大量附着生物,故本实验可排除其对水生植物的影响。本文证实了蓝藻死亡腐烂后的降解液极有可能是导致沉水植物消亡的主要原因。至于是由于蓝藻腐解液引起的哪种因素起主导作用,还有待进一步研究。     

砷形态对黑藻和竹叶眼子菜有机酸含量的影响

采用室内溶液培养法,运用气相色谱-质谱联用(Gas chromatography-mass spectrometry,GC-MS)技术测定在As(Ⅲ)、As(Ⅴ)和二甲基胂酸(Dimethylarsinic acid,DMA)3种形态砷(浓度0—5.0 mg/L)胁迫下,砷超富集植物黑藻(Hydrilla verticillata(Linn.f.)Royle.)和非砷超富集植物竹叶眼子菜(Potamogeton malaianus Miq.)体内生成和体外分泌的7种有机酸的含量。结果表明,黑藻体内草酸、丙二酸、柠檬酸、苹果酸和棕榈酸都显著高于竹叶眼子菜(P<0.05)。在As(Ⅲ)处理时,黑藻体内柠檬酸、苹果酸、棕榈酸、亚麻酸和总有机酸显著下降,竹叶眼子菜棕榈酸显著下降;在As(Ⅴ)处理时,黑藻棕榈酸和总有机酸显著下降,竹叶眼子菜棕榈酸显著下降;在DMA处理时,黑藻草酸、柠檬酸和苹果酸显著下降,竹叶眼子菜棕榈酸、亚麻酸和总有机酸显著下降。黑藻在As(Ⅲ)处理下棕榈酸、亚麻酸、总有机酸和As(Ⅴ)处理下草酸含量与体内As含量呈显著负相关(P<0.05),而竹叶眼子菜在As(Ⅲ)或DMA处理下棕榈酸、亚麻酸、总有机酸含量以及在DMA处理下柠檬酸含量与体内As含量均呈显著负相关(P<0.05)。两种植物根系分泌物中均检测出草酸、丙二酸、琥珀酸和棕榈酸,但只有在As(Ⅲ)或As(Ⅴ)处理时黑藻分泌的草酸与体内As含量呈显著正相关。本研究表明As(Ⅲ)或As(Ⅴ)胁迫下黑藻向体外分泌草酸是其超富集砷的一种重要机制。     

Light-dependent phosphate uptake of a submersed macrophyte Myriophyllum spicatum L.

The uptake kinetics of phosphate (Pi) by Myriophyllum spicatum was determined from adsorption and absorption under light and dark conditions. Pi uptake was light dependent and showed saturation following the Michaelis-Menten relation (in light: V = 16.91 × [Pi](1.335 + [Pi]), R² = 0.90, p < 0.001; in the dark: V = 5.13 × [Pi](0.351 + [Pi]), R² = 0.77, p < 0.001). Around 77% of the loss of Pi in the water column was absorbed into the tissue of M. spicatum, and only 23% was adsorbed on the surface of the plant shoots. Our study shows that M. spicatum shoots have a much higher affinity (in light: 3.9 μmol g⁻¹ dw h⁻¹ μM⁻¹; in the dark: 3.7 μmol g⁻¹ dw h⁻¹ μM⁻¹) and V_max (maximum uptake rate, shoot light) for Pi uptake than many other aquatic macrophytes (in light: 0.002-0.23 μmol g⁻¹ dw h⁻¹ μM⁻¹; in the dark: 0.002-0.19 μmol g⁻¹ dw h⁻¹ μM⁻¹), which may provide a competitive advantage over other macrophytes across a wide range of Pi concentrations.     

Growth,biomass allocation,and autofragmentation responses to root and shoot competition in Myriophyllum spicatum as a function of sediment nutrient supply

Plant growth, biomass allocation and autofragmentation were investigated in response to root and shoot competition in the submersed macrophyte Myriophyllum spicatum L. growing in two sediment environments. Biomass accumulation and allocation were significantly affected by sediment fertility, with a higher total biomass observed in fertile sediment (average: 4.69 g per plant vs. 1.12 g per plant in infertile sediment). Root-to-shoot ratios were 0.34 and 0.06 in the infertile and fertile sediments, respectively, reflecting the high investment placed on roots under infertile conditions. In the presence of root, shoot, and full competition, whole plant biomass decreased by 18%, 12% and 24% in the infertile sediments, and 23%, 25% and 33% in the fertile sediments, respectively. Root weight ratios (RWRs) increased with root competition by 38% (P < 0.001) and 12% (P = 0.002), while leaf weight ratios (LWRs) decreased with shoot competition by 6% (P = 0.042) and 5% (P = 0.001) in the infertile and fertile sediments, respectively. A total of 406 autofragments were harvested in the fertile sediments, but none were obtained from the infertile sediments. In the control, autofragment number and biomass was 166% and 175% higher compared to the competition treatment. Root and shoot competition resulted in a 21% (P = 0.043) and 18% (P = 0.098) decrease in the autofragment biomass, respectively. These results indicated that M. spicatum responds to different sediment fertility by changing its allocation patterns. Moreover, both root and shoot competition influenced plant growth and autofragmentation, while sediment nutrient availability played an important role in M. spicatum autofragmentation.     

Expansion of Myriophyllum spicatum (Eurasian water milfoil) into Lake Nasser,Egypt: Invasive capacity and habitat stability

The invasion of Myriophyllum spicatum into Lake Nasser, and its impact on submerged macrophyte communities are quantitatively documented. Samples of macrophytes, water and hydrosoil were collected from 17 sites, in October and November 2002. The average dry weight standing crop of each species per grapnel haul was determined at each depth zone (sampling site). Twenty-one environmental variables were measured (12 water and 9 hydrosoil variables). Canonical correspondence analysis (CCA) was used to determine species–environment relationships. Comparing these relationships of the present study with those detected in 1988–1990 indicated significant changes in water and hydrosoil characteristics. These changes are also implicated in the submerged macrophyte communities. M. spicatum has replaced the originally dominant submerged macrophyte Najas marina subsp. armata. The study indicated that the invasion of M. spicatum depends not only on its attributes, but also on the physico-chemical characteristics of Lake Nasser.     

Inter-annual variability of Potamogeton perfoliatus stands

Yearly differences in the composition and nutrient content of submerged macrophyte stands, mainly of the long-term dominant Potamogeton perfoliatus L. were observed at 10 stations along 83 km route of the Estonian shore of L. Peipsi (total area 3555 km²) in the period of 1999–2002. Changes consisted in the differences in the presence of plants (three stations), replacement of dominant (one station), removal of mud sediment (one station), mass propagation of Lemna trisulca (all stations) in 1999 and of Cladophora glomerata on P. perfoliatus (six stations) in 2000. The amount of C. glomerata on pondweed, similar to the biomass of host plant, decreased in 2001–2002, whereas markedly thick layer of epiphytic microalgae covered P. perfoliatus in 2002. In the samples of P. perfoliatus collected at the stations of the mass propagation of C. glomerata in the next year, 2001, inflorescences were absent. The appearance of an increase in P. perfoliatus stands was probably associated with improving sediment conditions in the period of low water level in the last observation years. A trend of constant increase in the mean molar N/P ratio in the tissues of P. perfoliatus in the study period (from 17.8 to 25.2) was statistically significant (p = 0.003). Classical canonical analysis, used to study two complexes correlated between themselves characteristics of P. perfoliatus and environmental conditions, revealed that plant growth factor (including biomass, tissue N%, P% and N/P) was in strong correlation (p < 0.0001) with water condition factor (including mean temperature in May, sum of temperatures between ice break and sampling, mean water level, N_tot and P_tot of water in June). More important for water condition factor were mean temperature in May and P_tot of water in June. Also the independent significance of water N_tot and P_tot for pondweeds was tested and confirmed by ANOVA analysis.     

Genetic diversity of an endangered aquatic plant, Potamogeton lucens subspecies sinicus

The genetic diversity of the species, Potamogeton lucens subsp. sinicus var. teganumensis, which is critically endangered in Japan, was investigated. This species now occurs in only two known localities in Japan. One is a native population (Oitoike population), but the other (Teganuma-Okahotto population) is found in a small artificial pond that was dug in 1998. It is considered that the Teganuma-Okahotto population grew from a soil seed bank. Based on RAPD variation, we compared the genetic diversity of the two populations of P. lucens var. teganumensis in Japan and one population of P. lucens subsp. sinicus var. sinicus in China. The Teganuma-Okahotto population showed RAPD variation, suggesting that it may be derived from more than one seed buried in old sediments. This population also had the highest value of Shannon's Information Index among the three study populations. This finding suggests that seeds buried in sediments can contain genetic variability, and may be used to conserve the genetic diversity of rare and endangered plants.     

Nuclear and chloroplast DNA sequences data support the origin of Potamogeton intortusifolius J.B. He in China as a hybrid between P. perfoliatus Linn. and P. wrightii Morong

Internal transcribed spacers (ITS) of nuclear ribosomal DNA and chloroplast rbcL gene sequence data were used to test the hypothesis that natural populations of Potamogeton intortusifolius J.B. He in China originated from hybridization between P. perfoliatus Linn. and P. wrightii Morong. Based on ITS sequences data, P. intortusifolius possessed heterozygous rDNA genotypes which confirmed the hybrid origin of P. intortusifolius. Chloroplast rbcL gene sequences of P. intortusifolius from Yichang population revealed the same chloroplast haplotype as P. perfoliatus and the samples of P. intortusifolius from Weinan population had the same chloroplast haplotype as P. wrightii, which indicated that both putative parental species had been the maternal parent and that the two populations of P. intortusifolius had independent origins. This study confirms P. intortusifolius as a reciprocal hybrid. Because P. × intortusifolius in China has the same hybrid origin as P. × anguillanus Koidz. in Japan, it is suggested that P. × intortusifolius should be a synonym of P. × anguillanus.     

菹草附着物对营养盐浓度的响应及其与菹草衰亡的关系

在菹草衰亡阶段对5个静水水体中菹草叶片表面附着物进行野外调查,并将其与水体营养盐浓度、沉水植物衰亡程度进行相关性分析。附着物共调查Chl.a含量、干重、有机质含量和藻类数量4个指标,沉水植物衰亡程度用单位面积叶片Chl.a含量表示,水体营养盐含量测量了TP、TN和N/P 3个指标。结果显示：附着物生物量与水体营养盐状况存在一定的正相关;各个点的附着物生物量与菹草衰亡状况存在一定相关性但相关性趋势与水体污染程度有关。在污染程度较高的水体中附着物生物量与菹草衰亡程度呈正相关,在污染程度较低的水体中附着物生物量与菹草衰亡程度呈负相关。结论为富营养化湖泊中营养盐含量的增加会导致附着物生物量的增加,但附着物只在污染程度较高的水体中促进植物衰亡。     

Cold stratification,light and high seed density enhance the germination of Ottelia alismoides

The effects of cold stratification, light and seed clustering in petri dish on Ottelia alismoides seed germination were investigated. The seeds required light and an extended cold period in order to germinate, but neither treatment alone was effective. Seed germination significantly increased with length of the 4 °C cold stratification period. Freshly collected seeds failed to germinate while a 5-month period at 4 °C yielded 29 ± 9% germination in the light, but none in the dark. Treatment with sodium nitroprusside, a nitric oxide source, failed to promote germination in the light or dark. Seeds of O. alismoides showed an unusual and significant positive response to aggregation. Germination in the light, after 5-month 4 °C cold stratification, was stimulated to almost five-fold in the dishes that were more densely sown with seed (20 seeds versus 200 seeds). Likewise, clustering seeds in dense aggregations stimulated germination significantly. Germination more than quadrupled with an increase from 1 to 50 seeds per cluster (200 seeds per dish), reaching a value of 72 ± 4%. Linear regression analysis shows the correlation between seed cluster density (no. per cluster) and germination rate (%) was highly significant (R² = 0.85, P = 0.000). The extended cold stratification requirement is probably an over-wintering device. The mechanism of the density-dependent stimulation is unclear.     

Seed germination of Lasia spinosa as a function of temperature,light, desiccation,and storage

Lasia spinosa seeds were not dormant at maturity in early spring. The most favorable temperatures for germination were between 25 and 30 °C, and final percentage and rate of germination decreased with an increase or decrease in temperature. When L. spinosa seeds were transferred to 25 °C, after 60 days at 10 °C (where none of the seeds germinated), final germination increased from 0% to 78%. Seeds germinated to high percentage both in light and in dark, although dark germination took more than twice as long as in the light. During desiccation of seeds at 15 °C and 45% relatively humidity, moisture loss decreased exponentially from 2.02 to 0.13 g H₂O g⁻¹ dry wt within 16 days, and only a few seeds (12%) survived 0.13 g H₂O g⁻¹ dry wt moisture content. Seeds stored at 0.58 g H₂O g⁻¹ dry wt moisture content at four constant temperatures (4, 10, 15, and −18 °C) for up to 6 months exhibited a well-defined trend of decreasing viability with decreasing temperature. Thus, we concluded that freshly harvested L. spinosa seeds are non-dormant and recalcitrant. Also, the seeds with 0.58 g H₂O g⁻¹ dry wt moisture content could be effectively stored for a few months between 10 and 15 °C although the most appropriate temperature for wet storage appears to be 10 °C, as it is close to the minimum temperature for germination and so there will be less pre-sprouting compared to 15 °C.     

Effects of pre-treatments and temperature on seed viability and germination of Juniperus macrocarpa Sm.

The effects of collecting season, collection site, laboratory pre-treatments and temperatures on seed viability and germination of Juniperus macrocarpa were investigated. Ripe cones were collected in four Sardinian dune systems, in two seasons, from plant and soil. Warm (W) and cold (C) stratifications, two combinations of them (W + C, C + W), and no pre-treatment (0) were applied. Seeds were incubated in a range of constant (10–25 °C) and an alternating (25/10 °C) temperature regime. Seed viability was low (ca. 40%) and varied significantly according to the collecting season. Seed germination was also low (ca. 10%), the 0 and W were the most effective pre-treatments on stimulating germination. The best germination temperature, without any pre-treatment, was 15 °C (ca. 20%). J. macrocarpa seeds are dormant and the achieved results suggested that the presence of secondary dormancy is induced by cold stratification. Spring appeared to be the best season for seed collecting, whereas autumn was the best for sowing. These results give new findings for restoration activities on this species.     

沙埋和种子大小对柠条锦鸡儿种子萌发、出苗和幼苗生长的影响

研究了沙埋深度和种子大小对内蒙古毛乌素沙地植被群落中占优势的柠条锦鸡儿(Caragana korshinskii Kom.)种子萌发、出苗、幼苗存活和生长的影响.结果表明,沙埋深度显著影响柠条锦鸡儿的种子萌发率、休眠率、出苗率、幼苗存活率及生物量.在0.5-2cm的浅层沙埋下,种子萌发率、出苗率、幼苗存活率及生物量最高,休眠率最低;沙埋深度≥4 cm时,柠条锦鸡儿的种子萌发率、出苗率、幼苗存活率及生物量随着沙埋深度增加显著降低,而休眠率却显著升高;沙埋深度≥12 cm时,柠条锦鸡儿种子不能够出苗,幼苗也不能够存活.种子大小对柠条锦鸡儿种子萌发率没有显著影响,但对出苗率、幼苗存活率及生物量影响显著.在各个沙埋深度下,不同大小的柠条锦鸡儿种子间的萌发率没有显著差异.当沙埋深度≤6 cm时,不同大小的柠条锦鸡儿种子在同一沙埋深度下的出苗率间没有显著差异;但当沙埋深度≥8 cm时,在同一沙埋深度下,大种子的出苗率显著高于中种子和小种子的出苗率,而中种子和小种子出苗率间没有显著差异.0.5-10 cm的沙埋深度中,除6 cm和8 cm深度下中种子和小种子萌发幼苗的生物量间没有显著差异外,其余深度下,大种子萌发的幼苗的存活率及生物量显著高于同一沙埋深度下中种子萌发的幼苗的存活率及生物量,后者又显著高于小种子萌发的幼苗的存活率及生物量.可能正是种子萌发对沙埋环境的忍耐或响应能力以及种子的多态性提高了柠条锦鸡儿在毛乌素沙地的适合度,为其在流动或半流动沙丘环境中成功定居并形成优势群落奠定了基础.     

Laboratory germination of seeds from 10 British species of Potamogeton

Factorial laboratory germination experiments were carried out on fruits from 10 British species of Potamogeton (Potamogetonaceae). Fruits had all been placed in a dry-room maintained at 15% RH and 15 °C for 3–6 months from the time of collection.     

A multi-assay comparison of seed germination inhibition by Lonicera maackii and co-occurring native shrubs

Recent work suggests that germination inhibition via allelopathy may be an important component of some species’ invasion ecology. We conducted four germination assays to test the inhibitory potential of Amur honeysuckle (Lonicera maackii), a problematic invasive shrub. The subject species in the first assay, Festuca arundinacea, exhibited a significant delay in germination when treated with extract from ground honeysuckle foliage. In the second assay, two concentrations of L. maackii extract were created by soaking foliage. F. arundinacea germination was not significantly influenced by the treatments while germination in Impatiens wallerana was substantially decreased. The third assay compared the impact of foliar extract from honeysuckle, with the native shrubs Lindera benzoin and Asimina triloba using F. arundinaceae, I. wallerana, Coreopsis lanceolata and Poa pratensis as subject species. In this assay, I. wallerana was strongly inhibited by L. maackii foliage; however, none of the other species were significantly influenced by the treatments. In the fourth assay, fruit extracts from L. maackii and L. benzoin were applied to the same four subject species. L. maackii fruit extract had an inhibitory influence on seed germination of all subject species, and inhibition from L. benzoin was also noted. Across all assays, there was a mixed reaction to extract from L. maackii and co-occurring native species that was species-specific and dependent upon the extract source. Our findings provide support for the idea that L. maackii has allelopathic activity, but further work is needed to (i) understand how broad the impact may be across the wide variety of species that are found in its invasion range and (ii) substantiate that the allelopathic effect has relevance in field environments.     

桔梗种子萌发对低温、干旱及互作胁迫的响应

以药用植物桔梗种子为材料,研究了低温、干旱及其互作对种子萌发和幼苗生长的影响,旨在探索在春寒干旱条件下其种子萌发对策及生态适应性,为桔梗野生种群恢复及人工栽培群体构建提供依据.结果表明,温度、干旱及其互作对桔梗种子萌发和幼苗生长均有显著影响.0-10％ PEG水分条件下,在变温10/20℃及恒温15-25℃下桔梗种子能够良好萌发(10％ PEG、20℃除外),而15％ PEG可显著抑制萌发.低温(10℃)下种子萌发始时间延迟,发芽率降低.随干旱胁迫程度的增加,10℃发芽率呈先升后降趋势,而其它温度下,发芽率和发芽指数均呈下降趋势.随温度的降低,幼苗根长、茎长及鲜重均呈下降趋势,且幼根生长对温度变化更为敏感.     

Effects of sand burial depth on seed germination and seedling emergence of Cirsium pitcheri

A greenhouse study was conducted to determine the effects of sand burial on seed germination and seedling emergence of Cirsium pitcheri, a threatened species along Lake Huron sand dunes. In October 1996, seeds of C. pitcheri were sorted into three groups (small, medium and large) and artificially buried at 2, 4, 6, 8, 10 and 12 cm depths in plastic pots filled with unsterilized sand from Lake Huron sand dunes. These pots were placed outside in sand boxes for natural stratification. In early February, the pots were brought into the greenhouse for germination of seeds. Data showed that percent seed germination and emergence of seedlings were not related to seed size. However, both variables were negatively correlated with depth. Seedling emergence occurred from a maximum depth of 6 cm with most seedlings emerging from 2 cm depth. Only one seedling emerged from 8 cm depth. Seedlings from large seeds produced longer roots than those of small seeds. Larger root system would probably enhance seedling establishment.