Do soil phytoliths accurately represent plant communities in a temperate region? A case study of Northeast China

Modern soil phytoliths can potentially provide analogues for phytolith assemblages from archaeological and palaeoecological contexts. To assess the reliability of soil phytoliths for representing different plant communities, we analysed phytoliths in surface soils and parent plants at 65 sites representing five types of regional vegetation in Northeast China. The results demonstrated that surface soil phytolith assemblages could clearly differentiate samples from herbaceous and woody communities, and samples from Poaceae and non-Poaceae communities could be separated statistically. In addition, woody communities could be differentiated into a broadleaf-Poaceae community, a broadleaf-non-Poaceae community and a conifer and broadleaf-non-Poaceae community, except for some overlapping samples. Soil phytolith assemblages are thus able to differentiate regional vegetation types into different plant community types. In the present study, soil phytoliths represented about 30% of the phytoliths present in the aboveground vegetation. In addition, soil phytoliths from different communities reflected the aboveground vegetation with slightly different degrees of accuracy, and in addition different morphotypes exhibited different degrees of representational bias. Some morphotypes (e.g. rondel, elongate psilate, lanceolate) overrepresented the abundance of the associated plant taxa; morphotypes such as tracheid, conical epidermal, stomata and others under-represented the original plant richness; and other morphotypes, e.g. saddle, trapeziform sinuate, scutiform, were in good agreement with the numbers of plant taxa in the plot inventory. Thus, any quantitative palaeovegetation reconstruction using phytoliths should begin with the calibration of soil phytolith assemblages. We conclude that our findings provide improved phytolith analogues for different plant communities, with applications in palaeoenvironmental reconstruction, and they also provide additional insights into the mechanisms of phytolith production and deposition.     

中国-中新世湖泊的古生态:根据山旺盆地植硅体的初步解释

山东山旺中新世山旺组湖相沉积中富含保存精美的古生物化石。对于湖相沉积中的昆虫、植物和哺乳动物化石,前人已经做了大量的研究工作。其中植物大化石和孢粉的研究结果表明,中新世山旺组在沉积期间,山旺湖周围的植被处于湿润的暖温带到亚热带的古生态环境下。哺乳动物蝙蝠和貘化石的研究结果也支持这个结论。然而,迄今为止,还没有关于草本植物的种类组成及含量的研究报道。本文首次对山旺组样品中的植硅体进行了研究,为山旺中新世植被的研究提供了新的证据。对从6块山旺组样品中分析出的植硅体进行研究的结果表明,山旺中新世植硅体组合含有丰富的保存极好的草本植物植硅体,代表森林成分的双子叶植物植硅体,以及偶尔出现的棕榈科植硅体。草本植物的植硅体主要来自于C_3/C_4禾本科的PACCAD类群和早熟禾亚科的植物,少数可能来自于在林下封闭生活环境中生长的草本植物。山旺中新世植硅体组合显示,在湖边森林生活环境下,伴生大量的适合沼生到中生环境的草本植物,在较干旱的地区生长着早熟禾(和PACCAD)的草本植物。本文的研究结果进一步支持山旺中新世气候比现在更加湿润而且变动较小的观点。山旺中新世植硅体组合与土耳其和北美大平原中新世湖相沉积中的植硅体组合不同,后者的植硅体组合主要是由多种C_3早熟禾亚科草本和不同种类的PACCAD类群草本植物组成。今后的研究工作将进一步揭示这种差异是否反映了草本植物群落在大尺度的生物地理上的差异,或者是由于地区性的局部气候不同所导致。     

内蒙古典型草原禾本科植硅体形态

运用地层中植硅体组合解释过去草原植被及气候变化的关键之一,是要了解研究区现代植硅体形态及表土植硅体组合与现代植被的关系。文中研究内蒙古典型草原禾本科植物根、茎、叶、芒以及种子等不同部位的植硅体,对其中的12种主要禾本科植物叶表皮短细胞硅酸体进行分类及统计。研究表明：内蒙古典型草原禾本科叶表皮短细胞硅酸体可分为8种特殊形态类型。C3植物早熟禾亚科的叶表皮短细胞硅酸体形态多样。几乎所有早熟禾亚科都能产生圆型硅酸体,以贝加尔针茅(85．5％)、大针茅(89．7％)、克氏针茅(90％)以及芨芨草(96．6％)中的圆型硅酸体含量最丰富。针茅哑铃型主要见于针茅植物叶表皮短细胞中,克氏针茅的针茅哑铃型含量相对较高。羊草中未见针茅哑铃型硅酸体。浴草、披缄草叶表皮短细胞硅酸体以齿型为主,分别含87．3％和57．2％,齿型在硬质早熟禾中也占一定比例。沙生冰草中的脊圆型占优势,含74．4％。C3植物早熟禾亚科的叶表皮短细胞产生的截锥型硅酸体含量较少。C4植物虎尾草亚科中的糙隐子草叶表皮短细胞硅酸体以黍哑铃型、简单哑铃型、鞍型为主;黍亚科狗尾草则以黍哑铃型占优势(82．9％)。     

Understanding Fossil Phytolith Preservation: The Role of Partial Dissolution in Paleoecology and Archaeology

Opaline phytoliths are important microfossils used for paleoecological and archaeological reconstructions that are primarily based on relative ratios of specific morphotypes. Recent studies have shown that phytolith assemblages are prone to post-depositional alteration involving partial dissolution, however, the manner in which partial dissolution affects morphotype composition is poorly understood. Here we show that morphotype assemblages from four different plant species subjected to controlled partial dissolution are significantly different from the original assemblages, indicating that the stability of various morphotypes differs, mainly depending on their surface area to bulk ratios. This underlying mechanism produces distorted morphotype compositions in partially dissolved phytolith assemblages, bearing vast implications for morphotype-based paleoecological and archaeological interpretation. Together with analyses of phytolith assemblages from a variety of archaeological sites, our results establish criteria by which well-preserved phytolith assemblages can be selected for accurate paleoecological and archaeological reconstructions.     

Phytolith assemblages in grasses native to central Argentina

BACKGROUND AND AIMS: Phytolith reference collections are a prerequisite for accurate interpretation of soil phytolith assemblages aimed at reconstructing past vegetation. In this study a phytolith reference collection has been developed for several grasses native to central Argentina: Poa ligularis, Piptochaetium napostaense, Stipa clarazii, Stipa tenuis, Stipa tenuissima, Stipa eriostachya, Stipa ambigua, Stipa brachychaeta, Pappophorum subbulbosum, Digitaria californica, Bothriochloa edwardsiana and Aristida subulata. METHODS: For each species, phytoliths present in the leaf blades were classified into 47 morphotypes, and their relative frequency determined by observing 300-400 phytoliths per sample (n = 5). Data were analyzed by complete linkage cluster analysis, using the Morisita Index as measure of association. KEY RESULTS: The results showed differentiation among phytolith assemblages at species level or at plant functional type level. Cluster analysis separated C3 from C4 species and palatable from non-palatable species. CONCLUSIONS: This study highlights the possibility of reconstructing past vegetation in central Argentina grasslands through the analysis of soil phytolith assemblages.     

Advantages and disadvantages of phytolith analysis for the reconstruction of Mediterranean vegetation: an assessment based on modern phytolith, pollen and botanical data (Luberon, France)

We present here the results of a first study comparing modern soil phytolith assemblages with pollen and botanical data at a North Mediterranean site. This work has shown the following limitations and advantages of phytolith analysis for the reconstruction of Mediterranean vegetation:(1) Phytoliths are produced in sufficient quantities for analysis and are well preserved in limestone environments, widespread in the Mediterranean area. (2) Young stands of Quercus ilex and Quercus coccifera, widely distributed in the Mediterranean area do not produce characteristic phytolith types in sufficient quantities to allow the calculation of a reliable index of tree cover density. (3) Pine forests, dominated by Pinus halepensis and Pinus sylvestris, are not recorded in the studied phytolith assemblages. (4) Grassland and shrubland assemblages can be distinguished through their associated phytolith assemblages, in particular by the proportion of crenate phytoliths produced in the short cells of the grass epidermis. (5) The different vegetation groups on the massif cannot be distinguished by pollen analysis. Further studies, comparing modern phytolith assemblages and quantitative vegetation data, should be carried out on forest plots that have been established for several centuries to further assess the role of phytolith analysis in vegetation reconstructions in the Mediterranean region.     

Assessing the importance of environmental factors to phytoliths of Phragmites communis in north-eastern China

Phytoliths have proved to be reliable indicators of environmental conditions both at present and in the past, and can provide evidence for the distribution of taxa or vegetation. Understanding the environmental significance of phytoliths within plants helps in drawing more reliable inferences about palaeovegetation and in reconstructing the palaeo-environment. The present study examined the relationship between phytoliths and environmental factors to assess the environmental significance of different types of phytoliths. Phytoliths were extracted from Phragmites communis growing in xerophytic and aquatic habitats at twelve sampling sites in north-eastern China and their implications for the environment were assessed using quantitative data mainly including phytolith concentration and environmental factors and statistical analyses. Principal component analysis (PCA) of several environmental factors (including the climate, micro terrain, and the physicochemical properties of soils) revealed that other factors being constant, P. communis phytolith concentrations were influenced largely by the average annual temperature and precipitation. Orthogonal experiment analysis and three-way analyses of variance of the concentrations confirmed the reliability of the results of the PCA. More specifically, the concentrations of the saddle and rondel types of phytoliths (the short-cell phytoliths) were closely linked to the average annual temperature whereas those of the elongate, lanceolate, and bulliform phytoliths (the non-short-cell phytoliths) were more sensitive to the variations in water status. The results contribute to further confirming the major environmental implications of phytoliths and, in turn, providing a reference for growth and development of wetland plants. Our findings also provide critical information that could help managers and policymakers assess and modify ecological restoration practices.     

Morphological variations of lobate phytoliths from grasses in China and the south-eastern United States

Abstract. Phytolith analysis of grasses is a useful tool in palaeoenvironmental and archaeobotanical research. Lobate phytolith is one of the most important morphotypes of grass phytoliths. This study describes morphological variations of diagnostic lobate phytoliths and produces a tentative classification scheme based on 250 modern grass species from China and the south‐eastern U.S.A. Eighty‐five grass species were found to contain lobate phytoliths. They are derived mainly from Panicoideae, but also include the Chloridoideae, Oryzoideae and Arundinoideae subfamilies. Twenty‐five lobate morphological types were observed from different subfamilies, genera or tribes of grasses, based on two important parameters: (1) the length of the lobate shank and (2) the shape of the outer margin of the two lobes. The identification of grass tribe or even genus is possible based on the differences in lobate shape parameters or the composition of assemblages. However, not all of the lobate assemblages have a definite relationship with the genera that produce them, because grasses can only produce a limited range of lobate shapes that often overlap from one genus to another. Several C₃ grasses and Chloridoideae subfamily grasses also produce characteristic lobate phytoliths. The variations of lobate morphologies can be related to environmental factors, especially moisture. Typical hygrophytic grasses tend to yield lobate phytoliths with very short shank, whereas typical xerophytic grasses tend to produce lobate phytoliths with a very long shank. The potential link between phytolith morphology, grass taxonomy and environmental conditions opens the possibility that phytolith morphology may be used as a proxy in palaeoclimatic reconstruction.     

New method for visualization of silica phytoliths in Sorghum bicolor roots by fluorescence microscopy revealed silicate concentration-dependent phytolith formation

Silica phytoliths are microscopic structures of amorphous hydrated silica (SiO₂·nH₂O) formed by specialized plant cells. Besides their biological roles, physical, chemical, and structural properties of biogenic silica offer a wide spectrum of applications in many fields of industry and technology. Therefore, processes involved in their formation recently become a very interesting topic to study. However, optical transparency and microscopic sizes of silica phytoliths do not allow their visualization and localization by classical light microscopy methods. Their observation thus requires phytolith isolation, technically difficult or lengthy sample preparation procedures, or a work with toxic chemicals. In this paper we are proposing a novel method for visualization of silica phytoliths in Sorghum bicolor root endodermal cells by fluorescence microscopy using alkali mounting solution (pH 12). This method offers an easy and quick preparation of the samples and high contrast imaging. Based on our results we can assume that the proposed fluorescent method for silica phytolith investigation allows observation of multiple samples in relatively short time period and thus might be applicable also for high-throughput screenings. Using this method we found out that after a 3-day cultivation of sorghum plants the minimal needed concentration of sodium silicate, limiting the formation of silica phytoliths in the root endodermis, was 25 µmol dm^?3. The positive correlation of sodium silicate concentration in the substrate with the phytolith diameter was also observed.     

Inter- and intraspecific variation in grass phytolith shape and size: a geometric morphometrics perspective

Background and AimsThe relative contributions of inter- and intraspecific variation to phytolith shape and size have only been investigated in a limited number of studies. However, a detailed understanding of phytolith variation patterns among populations or even within a single plant specimen is of key importance for the correct taxonomic identification of grass taxa in fossil samples and for the reconstruction of vegetation and environmental conditions in the past. In this study, we used geometric morphometric analysis for the quantification of different sources of phytolith shape and size variation.MethodsWe used landmark-based geometric morphometric methods for the analysis of phytolith shapes in two extant grass species (Brachypodium pinnatum and B. sylvaticum). For each species, 1200 phytoliths were analysed from 12 leaves originating from six plants growing in three populations. Phytolith shape and size data were subjected to multivariate Procrustes analysis of variance (ANOVA), multivariate regression, principal component analysis and linear discriminant analysis.Key ResultsInterspecific variation largely outweighed intraspecific variation with respect to phytolith shape. Individual phytolith shapes were classified with 83 % accuracy into their respective species. Conversely, variation in phytolith shapes within species but among populations, possibly related to environmental heterogeneity, was comparatively low.ConclusionsOur results imply that phytolith shape relatively closely corresponds to the taxonomic identity of closely related grass species. Moreover, our methodological approach, applied here in phytolith analysis for the first time, enabled the quantification and separation of variation that is not related to species discrimination. Our findings strengthen the role of grass phytoliths in the reconstruction of past vegetation dynamics.     

Phytoliths Analysis for the Discrimination of Foxtail Millet (Setaria italica) and Common Millet (Panicum miliaceum)

Foxtail millet (Setaria italica) and Common millet (Panicum miliaceum) are the oldest domesticated dry farming crops in Eurasia. Identifying these two millets in the archaeobotanical remains are still problematic, especially because the millet grains preserve only when charred. Phytoliths analysis provides a viable method for identifying this important crop. However, to date, the identification of millet phytoliths has been questionable, because very little study has been done on their morphometry and taxonomy. Particularly, no clear diagnostic feature has been used to distinguish between Foxtail millet and Common millet. Here we examined the anatomy and silicon structure patterns in the glumes, lemmas, and paleas from the inflorescence bracts in 27 modern plants of Foxtail millet, Common millet, and closely related grasses, using light microscopy with phase-contrast and microscopic interferometer. Our research shows that five key diagnostic characteristics in phytolith morphology can be used to distinguish Foxtail millet from Common millet based on the presence of cross-shaped type, regularly arranged papillae, Ω-undulated type, endings structures of epidermal long cell, and surface ridgy line sculpture in the former species. We have established identification criteria that, when used together, give the only reliable way of distinguishing between Foxtail millet and Common millet species based on their phytoliths characteristics, thus making a methodological contribution to phytolith research. Our findings also have important implications in the fields of plant taxonomy, agricultural archaeology, and the culture history of ancient civilizations.     

温带地区表土植硅体对温度的响应

植硅体的现代过程研究是利用植硅体这一指标精准恢复区域古植被、古气候的前提和关键环节,探讨表土植硅体组合的空间分布特征及其对环境因子的响应尤为重要.本研究在东北地区沿着年降水量为600 mm等降水线采集54块表土样品,研究单一温度控制下表土植硅体的空间分布规律,以期寻找对温度比较敏感的植硅体类型.结果表明:54块表土样品...     

东北地区芦苇植硅体的变化特征

深入研究芦苇(Phragmites australis)不同生长期植硅体的形态变化规律, 对提高古植被重建精度、探讨植硅体形成机理具有重要意义。该文对东北地区12个样点的芦苇叶片进行研究, 选取长势相近、叶片大小一致的芦苇叶片, 用湿式灰化法提取植硅体并对其进行分类和命名。研究发现, 芦苇植硅体主要有5种类型, 在不同生长期内和不同温湿度组合下, 其植硅体类型组合基本无变化, 说明同种植物的植硅体类型组合具有稳定性, 据此可以有效地恢复古植被; 同时芦苇鞍型植硅体百分含量的峰值出现在7月份, 随后稍有减少, 而芦苇扇型植硅体百分含量和浓度的峰值出现在8或9月份, 谷值在7月份。由此推测芦苇鞍型植硅体的形成可能更多地受控于植物的光合作用和蒸腾作用, 芦苇扇型植硅体的形成可能与机动细胞的支撑作用关系密切。实验结果还表明芦苇植硅体浓度并不是随着时间逐渐积累的, 其在8月份达到峰值, 9月份达到谷值, 因此推测植物植硅体浓度的变化规律可能与植物在不同生长期其自身对硅的需求规律相一致。鞍型植硅体浓度的峰值也出现在8月份, 谷值也在9月份, 其浓度变化与芦苇植硅体总浓度的变化趋势相似。     

东北地区芦苇植硅体形态的空间差异

笔者对东北地区12个样点的芦苇叶片的植硅体进行了分析,探讨了芦苇植硅体在空间上的分布规律,揭示芦苇植硅体的环境指示意义,为定量恢复古环境提供参考。实验结果表明:12个样点中芦苇植硅体基本类型相同,共有5种主要植硅体类型,即中鞍型、帽型、芦苇扇型、尖型和棒型;但芦苇植硅体的浓度随着纬度的变化而相应的发生变化。总体来说,纬度较低的样点芦苇植硅体的浓度较大,而纬度较高的样点芦苇植硅体的浓度相对较小;随着纬度的变化,芦苇的特征植硅体中鞍型的浓度也呈现波状变化,牡丹江以南的几个样点中鞍型植硅体浓度相对较大,9、10两个月牡丹江以北的几个样点中鞍型植硅体浓度迅速减少。芦苇扇型植硅体则呈现出纬度较高的样点植硅体浓度较大,纬度较低的样点植硅体浓度较小的特点,芦苇扇型植硅体的空间变化规律与扇型植硅体的不同。     

Disease Interactions in a Shared Host Plant: Effects of Pre-Existing Viral Infection on Cucurbit Plant Defense Responses and Resistance to Bacterial Wilt Disease

Both biotic and abiotic stressors can elicit broad-spectrum plant resistance against subsequent pathogen challenges. However, we currently have little understanding of how such effects influence broader aspects of disease ecology and epidemiology in natural environments where plants interact with multiple antagonists simultaneously. In previous work, we have shown that healthy wild gourd plants (Cucurbita pepo ssp. texana) contract a fatal bacterial wilt infection (caused by Erwinia tracheiphila) at significantly higher rates than plants infected with Zucchini yellow mosaic virus (ZYMV). We recently reported evidence that this pattern is explained, at least in part, by reduced visitation of ZYMV-infected plants by the cucumber beetle vectors of E. tracheiphila. Here we examine whether ZYMV-infection may also directly elicit plant resistance to subsequent E. tracheiphila infection. In laboratory studies, we assayed the induction of key phytohormones (SA and JA) in single and mixed infections of these pathogens, as well as in response to the feeding of A. vittatum cucumber beetles on healthy and infected plants. We also tracked the incidence and progression of wilt disease symptoms in plants with prior ZYMV infections. Our results indicate that ZYMV-infection slightly delays the progression of wilt symptoms, but does not significantly reduce E. tracheiphila infection success. This observation supports the hypothesis that reduced rates of wilt disease in ZYMV-infected plants reflect reduced visitation by beetle vectors. We also documented consistently strong SA responses to ZYMV infection, but limited responses to E. tracheiphila in the absence of ZYMV, suggesting that the latter pathogen may effectively evade or suppress plant defenses, although we observed no evidence of antagonistic cross-talk between SA and JA signaling pathways. We did, however, document effects of E. tracheiphila on induced responses to herbivory that may influence host-plant quality for (and hence pathogen acquisition by) cucumber beetles.     

Epidermal Patterning and Silica Phytoliths in Grasses: An Evolutionary History

Due to the immense ecological and economic significance of grasses, their highly characteristic long–short epidermal patterning and associated silica phytoliths represent significant diagnostic markers in studies of ancient climate change and agriculture. We explore the link between epidermal cell patterning and phytolith development and review the evolutionary history of phytoliths in the context of recent well-resolved phylogenetic analyses of grasses and allied Poales, focusing on early-divergent grasses and the subfamilies that constitute the BEP group (the bamboos and their allies). Dimorphic epidermal patterning is a common feature of Poaceae and the related family Joinvilleaceae, where phytoliths are located primarily in the short cells. However, Joinvillea lacks the short-cell pairs that occur in many grasses. The costal rows of phytoliths that characterize some grasses could represent loss of long–short cell patterning over the veins. Unlobed phytoliths probably represent the ancestral condition in grasses, though bilobate phytoliths evolved at an early stage. Either transverse-unlobed or transverse-bilobate phytoliths predominate in the early-divergent lineages, whereas axial-bilobates (or polylobates) primarily characterize the PACMAD clade and the BEP subfamily Pooideae.     

Do phytoliths play an antiherbivory role in southwest Asian Asteraceae species and to what extent?

Phytoliths (silica bodies) occur in Poaceae species in large numbers and have been shown to have antiherbivory roles. However, phytoliths occur also in many other taxa in much smaller numbers, which raises the question of the extent of both their potential and actual antiherbivory role in these taxa. In order to address the question of their potential antiherbivory role, we sampled 20 wild-growing southwest Asian species of the family Asteraceae, species of which have a much lower phytolith concentrations than Poaceae taxa. We studied the potential positive effect of grazing on phytolith formation and the possible tendency of plants to have higher concentrations of such defence structures in their reproductive organs. We sampled plants from populations of 12 non-spiny and eight spiny species growing in un-grazed and grazed plots in seven sites along a large rainfall gradient (80–900 mm mean annual) in Israel, a region known for its long and intensive grazing history. The study included 21 pairs of un-grazed and grazed plants from 16 of these 20 species. In addition, ten populations of eight species were sampled in order to examine whether phytolith concentrations in the reproductive organs are higher than in vegetative organs. We did not find consistently higher phytolith concentrations in grazed plants compared to un-grazed plants of the same species and habitat (15 species), and in 12 out of 21 pairs of un-grazed and grazed plants (from 15 species) we even found higher phytolith concentrations in un-grazed plants, a phenomenon which was more common in the more arid sites. Phytolith concentrations in inflorescences are commonly (6 out of the 8 species) lower than in the rest of the shoot. We conclude that the antiherbivory potential of phytoliths in the southwest Asian Asteraceae as a group is much smaller than in the Poaceae.     

Assessing environmental and plant community changes by phytolith characteristics in sediments from Nanjishan Nature Reserve,Lake Poyang

To assess the impacts of increasing human disturbance, wetland degradation, and biodiversity loss, and to determine the main environmental factors affecting the spatial distribution of phytoliths in Nanjishan Nature Reserve, phytoliths from 17 topsoil samples and a sediment core sample were analyzed. Yellow-brown, silty, and gray-black clays were the main sedimentary constituents, and the total organic carbon content decreased with increasing depth. Lakes Zhanbei, Chang, and Sanniwan were located in an alluvial delta and characterized by intensive human activity and grass communities of Triarrhena lutarioriparia and Zizania latifolia, whereas Lakes Shangbeijia, Beishen, Nanshen and Baisha were characterized by a lower degree of disturbance and grass communities of T. lutarioriparia and Phragmites australis. A total of 11,942 phytoliths (4248 in topsoil and 7694 in core sediment) were isolated from 46 samples and 19 distinct morphological types were identified. Vesicular cells constituted 42%–51% of all samples, short cells 24%–32%, and elongate types 15%–26%. The phytolith assemblages at Lakes Zhanbei, Chang, and Sanniwan were mainly composed of fans, rectangles, points and elongates. While Lake Shangbeijia, Beishen, Nanshen and Baisha were dominated by fans, saddles and elongates. Based on phytolith analysis, we found that the typical communities in the study area were significantly degraded from 1986 to 2003. Sediment cores dated by ²¹⁰Pb showed that the main plant communities were transformed from Carex and T. lutarioriparia to Z. latifolia, T. lutarioriparia and P. australis.