Advancing the use of minirhizotrons in wetlands

Background

Wetlands store a substantial amount of carbon (C) in deep soil organic matter deposits, and play an important role in global fluxes of carbon dioxide and methane. Fine roots (i.e., ephemeral roots that are active in water and nutrient uptake) are recognized as important components of biogeochemical cycles in nutrient-limited wetland ecosystems. However, quantification of fine-root dynamics in wetlands has generally been limited to destructive approaches, possibly because of methodological difficulties associated with the unique environmental, soil, and plant community characteristics of these systems. Non-destructive minirhizotron technology has rarely been used in wetland ecosystems.

Scope

Our goal was to develop a consensus on, and a methodological framework for, the appropriate installation and use of minirhizotron technology in wetland ecosystems. Here, we discuss a number of potential solutions for the challenges associated with the deployment of minirhizotron technology in wetlands, including minirhizotron installation and anchorage, capture and analysis of minirhizotron images, and upscaling of minirhizotron data for analysis of biogeochemical pools and parameterization of land surface models.

Conclusions

The appropriate use of minirhizotron technology to examine relatively understudied fine-root dynamics in wetlands will advance our knowledge of ecosystem C and nutrient cycling in these globally important ecosystems.     

Spatial variation in tree root distribution and growth associated with minirhizotrons

Four minirhizotrons were installed in each of three replicate plots in a deciduous forest dominated by Acer saccharum Marsh. The length growth of tree roots along the surface of the minirhizotrons was measured for a period of one year, and the resulting data were analyzed in nested, averaged and pooled arrangements. The analyses of nested data showed that spatial variation in root growth and abundance among minirhizotrons within plots was greater than variation among plots. Averaging data from minirhizotrons within plots prior to analysis reduced variation about plot means, but extensive intraplot variation invalidates this approach on statistical grounds. Both nested and averaged data failed to account for the contribution of individual roots to the mean, and root production rates were consequently overestimated. Pooling the data from the four minirhizotrons reduced variation about the means, and resulted in a more representative estimate of root production rates. The analysis of composited data can be used to incorporate small-scale variation into a single replicate sample in those circumstances where the activity of the root systems of plant communities is the object of study.     

Applications and limitations of rhizotrons and minirhizotrons for root studies

This paper discusses two glass-wall techniques-rhizotrons and minirhizotrons. Rhizotrons are covered underground walkways with clear windows on one or both sides. Their design varies somewhat with the type of research to be conducted in them. A minirhizotron consists of a clear tube inserted into the soil then some type of viewing arrangement is lowered into the tube. In both techniques, roots and their rhizosphere organisms can be observed at the soil-wall boundary.     

Applications of minirhizotrons to understand root function in forests and other natural ecosystems

Minirhizotrons have proved useful to understand the dynamics and function of fine roots. However, they have been used comparatively infrequently in forests and other natural plant communities. Several factors have contributed to this situation, including anomalous root distributions along the minirhizotron surface and the difficulty of extracting data from minirhizotron images. Technical and methodological advances have ameliorated some of these difficulties, and minirhizotrons have considerable potential to address some questions of long standing interest. These questions include more fully understanding the role of roots in carbon and nutrient cycling, rates of root decomposition, responses to resource availability and the functional significance of interactions between plant roots and soil organisms. Maximizing the potential for minirhizotrons to help us better understand the functional importance of fine roots in natural plant communities depends upon using them to answer only those questions appropriate to both their inherent strengths and limitations.     

A new dimension to observations in minirhizotrons: A stereoscopic view on root photographs

Summary With a stereoscope, as used for the inspection of aerial photographs, sequential photographs of roots obtained by the endoscope method from minirhizotrons can yield much more information than hitherto. A series of photographs shows that most of the roots seen in a minirhizotron in grassland grew on the surface of the lexan tube, while there was a gap between the roots and the soil. Decay of the extensive root hair zones around the roots may make new root growth in the gap between rhizotron wall and soil invisible. Some consequences of these observations for the endoscope method are discussed.     

A comparison of soil core sampling and minirhizotrons to quantify root development of field-grown potatoes

Root growth of potato (Solanum tuberosum L.) is sensitive to soil conditions. A reduced root system size can result in reduced uptake of water and/or nutrients, leading to impaired crop growth. To understand the mechanisms by which soil conditions affect crop growth, study of temporal and spatial development of roots is required.In field experiments, effects of soil temperature, soil compaction and potato cyst nematodes (Globodera pallida) on root growth of potato cultivars were studied using two methods: core sampling and vertically oriented minirhizotrons.Minirhizotrons showed relatively more roots in deeper soil layers than core sampling, probably because of preferential root growth along the tube. Spatial distribution of roots should therefore be analysed by core sampling.To eliminate differences in spatial distribution, total root systems as measured by both methods were compared. Nematodes, cultivars and time did not affect the relationship between both methods. Soil compaction, however, affected it because of a strong response of root length in bulk soil and small differences in root number against the minirhizotron, suggesting that soil coring has to be used to study effects of different bulk densities.With both methods, sequential measurements of roots give the net effect of root growth and decay. Data on root turnover can only be obtained with minirhizotrons by comparing video recordings of different dates. Other information obtained with minirhizotrons is the average orientation of roots. Moreover, the minirhizotron method has the advantage of demanding less labour.     

Advancing realism in biodiversity research

Numerous experimental studies suggest that biodiversity loss is detrimental to ecosystem functions such as production and nutrient cycling. These experiments, however, have been criticized as unrealistic because they use combinations of species that do not resemble what is observed in nature. Bracken et al. take a novel approach, using species combinations observed in the field and computer simulations to explore many other combinations. Their approach represents a significant advance in making biodiversity research more realistic.     

Advancing research on ectomycorrhizal fungal adaptation with landscape genomics

Ectomycorrhizal (ECM) fungi serve key functions in forest ecosystems by supplying water and nutrients to tree hosts, yet mutualistic plant–fungi interactions are jeopardised by environmental alterations. Here, we discuss the great potential and current limitations of landscape genomics in investigating signatures of local adaptation in natural populations of ECM fungi.     

杨树人工林根际古菌群落随细根生长的演变

研究细根不同生长时期根际土壤古菌群落组成结构差异,对深入了解林木细根与土壤微生物互作关系具有重要理论意义.依据细根表面颜色,采集杨树一级细根不同生长时期(白色新生根、黄色成熟根、褐色衰老根)根际土壤并提取微生物总DNA,采用特异性引物对古菌16S rDNA V4-V5区进行扩增,利用Illumina MiSeq平台进行古菌高通量测序分析.结果表明: 新生根和衰老根根际土壤古菌群落操作分类单元(OTU)丰富度相似,而成熟根根际土壤古菌群落OTU数量较少.新生根和成熟根根际土壤共同含有134个OTU;成熟根和衰老根根际土壤共同含有87个OTU,新生根和衰老根根际土壤共同拥有90个OTU.α多样性分析表明,成熟根根际土壤古菌群落Chao1指数和ACE指数显著低于新生根和衰老根根际土壤,而衰老根根际土壤古菌群落Simpson指数和Shannon指数显著低于新生根和成熟根根际土壤.PERMANOVA分析表明,新生根和衰老根根际土壤古菌群落组成有显著差异.物种注释显示,杨树根际土壤共包含12个古菌属,其中新生根5个、成熟根10个、衰老根6个.β多样性指数表明,杨树根际土壤古菌群落相似度随着细根的生长逐渐下降,不同生长阶段细根根际土壤的古菌群落结构有较大差异.其中,占绝对优势的古菌为氨氧化古菌Candidatus_Nitrososphaera,其相对丰度超过70%.且随细根生长发育,该类古菌在根际土壤中的丰度呈现上升趋势,表明其可能与细根的生长发育关系密切.     

微根管在细根研究中的应用

细根（直径≤2 mm）的周转在植物生态系统碳分配过程中具有重要意义.已往细根周转研究主要采用根钻法、分室模型法和内生长法等.这些方法由于不能直接观测到细根生长动态,导致细根周转估计不准确.微根管法是一种非破坏性野外观察细根动态的方法.本文从微根管的发展、功能、安装步骤、图像采集、参数计算、影响观测因素和存在问题等方面逐一进行介绍,并通过水曲柳和落叶松微根管细根观测实例介绍在细根周转过程研究中的应用. 结果表明,微根管可以比较精确地估计出细根长度、单位面积上根长密度、单位体积上根长密度、细根生长量、细根死亡量和细根周转等.微根管是一个观察细根生长、衰老、死亡和分解过程的有效工具.微根管观测精度主要取决于微根管安装的质量和数量、微根管取样间隔期和取样数量、微根管图像分析技术等.此外,土壤质地、石砾多少、微根管材料选择、减少光系统对根系的干扰等也是影响微根管测定精度的因素.如何提高微根管测定精度将成为今后微根管在细根研究中的主要问题.     

Stand fine root biomass and fine root morphology in old-growth beech forests as a function of precipitation and soil fertility

Only very limited information exists on the plasticity in size and structure of fine root systems, and fine root morphology of mature trees as a function of environmental variation. Six northwest German old-growth beech forests (Fagus sylvatica L.) differing in precipitation (520 – 1030 mm year^–1) and soil acidity/fertility (acidic infertile to basic fertile) were studied by soil coring for stand totals of fine root biomass (0–40 cm plus organic horizons), vertical and horizontal root distribution patterns, the fine root necromass/biomass ratio, and fine root morphology (root specific surface area, root tip frequency, and degree of mycorrhizal infection). Stand total of fine root biomass, and vertical and horizontal fine root distribution patterns were similar in beech stands on acidic infertile and basic fertile soils. In five of six stands, stand fine root biomass ranged between 320 and 470 g m^–2; fine root density showed an exponential decrease with soil depth in all profiles irrespective of soil type. An exceptionally small stand fine root biomass (<150 g m^–2) was found in the driest stand with 520 mm year^–1 of rainfall. In all stands, fine root morphological parameters changed markedly from the topsoil to the lower profile; differences in fine root morphology among the six stands, however, were remarkably small. Two parameters, the necromass/biomass ratio and fine root tip density (tips per soil volume), however, were both much higher in acidic than basic soils. We conclude that variation in soil acidity and fertility only weakly influences fine root system size and morphology of F. sylvatica, but affects root system structure and, probably, fine root mortality. It is hypothesized that high root tip densities in acidic infertile soils compensate for low nutrient supply rates, and large necromasses are a consequence of adverse soil chemical conditions. Data from a literature survey support the view that rainfall is another major environmental factor that influences the stand fine root biomass of F. sylvatica.     

Consequences of insect herbivory on grape fine root systems with different growth rates

Herbivory tolerance has been linked to plant growth rate where plants with fast growth rates are hypothesized to be more tolerant of herbivory than slower-growing plants. Evidence supporting this theory has been taken primarily from observations of aboveground organs but rarely from roots. Grapevines differing in overall rates of new root production, were studied in Napa Valley, California over two growing seasons in an established vineyard infested with the sucking insect, grape phylloxera (Daktulosphaira vitifoliae Fitch). The experimental vineyard allowed for the comparison of two root systems that differed in rates of new root tip production (a 'fast grower', Vitis berlandieri x Vitis rupestris cv. 1103P, and a slower-growing stock, Vitis riparia x Vitis rupestris cv. 101-14 Mgt). Each root system was grafted with a genetically identical shoot system (Vitis vinifera cv. Merlot). Using minirhizotrons, we did not observe any evidence of spatial or temporal avoidance of insect populations by root growth. Insect infestations were abundant throughout the soil profile, and seasonal peaks in phylloxera populations generally closely followed peaks in new root production. Our data supported the hypothesis that insect infestation was proportional to the number of growing tips, as indicated by similar per cent infestation in spite of a threefold difference in root tip production. In addition, infested roots of the fast-growing rootstock exhibited somewhat shorter median lifespans (60 d) than the slower-growing rootstock (85 d). Lifespans of uninfested roots were similar for the two rootstocks (200 d). As a consequence of greater root mortality of younger roots, infested root populations in the fast-growing rootstock had an older age structure. While there does not seem to be a trade-off between potential growth rate and relative rate of root infestation in these cultivars, our study indicates that a fast-growing root system may more readily shed infested roots that are presumably less effective in water and nutrient uptake. Thus, differences in root tip production may be linked to differences in the way plants cope with roots that are infested by sucking insects.     

施肥对日本落叶松人工林细根直径、根长和比根长的影响

以辽宁东部山区16年生日本落叶松人工林为研究对象,探讨施肥对日本落叶松1～5级根序中细根直径、根长和比根长的影响.结果表明：随着根序等级的增加,日本落叶松细根平均直径和根长显著增加（P<0。05,P<0。01）、比根长则显著下降（P<0。01）.在日本落叶松的5级根序中,1级根的平均直径最细、根长最短、比根长最高,而5级根则相反;随着根序等级的增加,日本落叶松细根平均直径、根长和比根长的变异系数逐渐增大.除1级根外,土层对细根的平均直径、根长和比根长没有显著影响(P>0。05).与对照样地相比,施肥对各级细根平均直径、根长和比根长的影响主要表现在1～2级根上,对3级根序以上的细根影响不显著（P>0.05）.其中,施氮肥显著降低了1～2级根的平均直径（P<0.05）,施氮肥以及氮磷肥显著降低了表层土壤（0～10 cm）中1级根的平均根长（P<0.05）,表层土壤中细根的比根长在施氮肥的条件下显著增加（P<0.05）.     

Reduced root anchorage of freshwater plants in sandy sediments enriched with fine organic matter

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