A Review on Diagnostic Phytoliths for the Application in Paleovegetation Reconstruction and Environmental Archaeology in East Asia

Phytoliths are extensively utilized as an archaeobotanical indicator in paleovegetation reconstruction and environmental archaeology. Over the past two decades, numerous phytolith morphotypes, particularly those exhibiting diagnostic morphological features and over representative of source plants at the genus and species levels, were discovered and reported. These advancements have significantly contributed to phytolith-based vegetation reconstruction on different timescales, enhanced our understanding of prehistoric plant utilization, and elucidated cultivation and domestication processes of key crops in ancient agriculture. However, there are still inconsistencies and misunderstandings regarding the morphological characteristics of diagnostic phytoliths in various plant groups. This review highlighted the standardization in the classification and description of phytolith morphotypes, and summarized the advancements in phytolith morphology research over the past two decades. Morphological illustrations of diagnostic phytoliths from various plant groups, particularly key crops and their relatives from dryland and rice agriculture in East Asia, were presented as references for phytolith identification and application. Finally, this review proposes future directions for phytolith morphological studies, emphasizing the comprehensive consideration of anatomical structure and morphometric parameters, as well as the need for extensive research on modern plant phytoliths and control experiments on phytolith growth.

对phytolith汉译名的商榷

基于对ｐｈｙｔｏｌｉｔｈ一词内涵和使用沿革的回顾，笔者主张将其译为植结石，可以与动结石（ｚｏｏｌｉｔｈ）匹对，不主张将其译为“植物硅酸体”、“植物硅石”、“植物硅体细胞”和“植物石”。同样，一门新兴的边缘学科──ｐｈｙｔｏｌｉｔｈｉｃｓ可译为植结石学。

国际植硅体命名法规(International Code for Phytolith Nomenclature 1.0)的介绍与讨论

随着植硅体分析的广泛应用,植硅体命名规范化已经成为趋势。文章对国际植硅体命名法规中植硅体类型划分、命名和描述的方法与术语进行介绍,提供部分中文对应译名,并对该法规中存在的问题进行探讨,以期引起国内学术界的重视,为我国植硅体命名规范化提供参考。

Pollen and Phytolith Analyses of Ancient Paddy Fields at Chuodun Site, the Yangtze River Delta

A number of paddy fields pertaining to the Majiabang Cultures (5500-3800 years BC) were discovered during the archaeological excavations that were carried out since 1998 at the Chuodun site in the Yangtze River Delta. The pollen and phytolith analyses of two soil profiles from the northeastern part of this site were carried out to trace the agricultural practices of the Neolithic period. The phytolith results showed that rice domestication in the Yangtze River Delta could be traced back to as early as the Majiabang Culture. The pollen assemblage also revealed low levels of aquatic species, similar to that in modern paddy fields. This finding suggested that humans might have removed weeds for rice cultivation during the Neolithic period. Thus, pollen analysis in association with phytolith analysis was a promising method for identifying ancient paddy fields.

Paleo-environmental evolution based on high-resolution phytolith since 26 ka B. P. in Huanghe River Delta

The Huanghe River Delta is one of the world＇s large rivers, the Huanghe River Delta paleoenvironmental evolution in the Huanghe River has been a hot issue since the Last Glacial. Based on the core time series established by combining AMS 14C dating of Core DYZK1 sediments in submerged of Huanghe River Delta and acoustic sequence on sub-bottom profile, phytolith analyses are carried out on 96 sediment samples. The grain size parameters, magnetic susceptibility are combined with the vertical changes of biostratum to reconstruct the paleo-sedimentary and climatic conditions in the Huanghe River Delta. The study results show that there is a significant vertical change law in the index parameters, and that sedimentary environment of Huanghe River Delta experienced an evolutionary process of fluvial facies-sealand transition facies-tidal flat facies-neritic facies- delta facies since 26.0 ka B. P.. The phytolith analyses results are as following. Firstly, The phytoliths are divided into lanceolate, smooth-elongated, spiny-elongated, rondel, long rectangular, bulliform and other types. In different sedimental environment, the phytolith content changes regularly, indicating that the phytolith assembles in the same sedimentary environment has a certain degree of stability. Secondly, The lanceolate, smooth-elongated and spiny-elongated categories of phytoliths have greater contents in the tidal flat facies and delta deposition, while have a smaller contents in the neritic facies and fluvial facies environment. Thirdly, Through comparative analysis of variation coefficient, the content of major phytolith categories in the sediment has a greater change amplitude in the sealand transition facies and fluvial facies deposition, while being more stable in the tidalflat facies and delta facies deposition.

Contribution of phytoliths to total biogenic silica volumes in the tropical rivers of Malaysia and associated implications for the marine biogeochemical cycle

The contribution of phytoliths to total biogenic silica(BSi) volumes in rivers worldwide,and the associated implications for the biogeochemical cycle,require in-depth study.Based on samples from rivers in Peninsular Malaysia,this project investigated the source and characteristics of B Si found in Asian tropical rivers,as well as the process of reverse weathering taking place in these fluvial systems.Results indicated that BSi samples collected in sediments consisted of phytolith,diatom and sponge spicules.Phytoliths,predominantly of the elongate form,comprised 92.8%-98.3% of BSi in the Pahang River.Diatom BSi in this river consisted mainly of pennatae diatoms,but represented a relatively small proportion of the total BSi volume.However,diatom BSi(predominantly of the Centricae form) was more prevalent in the Pontian and Endau Rivers with shares of 68.8% and 79.3% of the total BSi volumes,respectively,than Pahang River.Carbon contents of the BSi particulates ranged from 1.85% to 10.8% with an average of 4.79%.These values are higher than those recorded in other studies to date,and indicate that BSi plays a major role in controlling permanent carbon burial.This study suggests that phytoliths from terrestrial plants are the primary constituents of BSi in the rivers of Peninsular Malaysia,and therefore represent a significant proportion of the coastal silica budget.

Responses of Phytolith in Guinea Grass(Leymus chinensis) Leaves to Simulated Warming,Nitrogen Deposition and Elevated CO_2 Concentration in Songnen Grassland,Northeast China

Deposited in plant cells and their intercellular space,phytoliths,a special form of silica,could be used to determine information on plant structure and physiology especially their size and content.With the hypothesis that phytolith in plant would change under variable climate and environment,the dominant plant species in Songnen grassland,guinea grass(Leymus chinensis),was treated by an open-top chamber(OTC) to elevate CO2 concentration,infrared heaters,and artificial nitrogen(N) addition for three years from 2006–2008.Phytoliths were extracted by wet-ashing method and analyzed by variance analysis and so on.We found that the responses to elevated CO2 are complicated,and warming is positive while N addition is negative to the deposition of phytoliths in L.chinensis leaves.Especially,warming could reduce the negative impact of N addition on phytolith in L.chinensis.The short cell's taxonomic in graminea is significant because of no disappearance with simulated environmental changes.The phytolith originated in the long cell and plant intercellular space are more sensitive to elevated CO2 concentration,warming,and N addition,and could become some new indicators for environmental changes.In conclusion,different phytolith types have various responses to simulated warming,N addition and elevated CO2 concentration.

The Need of Time:Phytoliths for Earth Science

Phytoliths take the shape and size of the cells from which they are formed.They are microscopic and composed of opal silica—SiO2 and other traces of Al, Fe,Mn,P,Cu,N and organic carbon.Phytoliths are present in the roots,leaves,stem and the rest parts of the plant body including inflorescences.The shapes of phytolith can be used to determine the taxonomy of the plants,which may be dumbbell,squares,rectangular,

The Phytolith and Pollen Record since 10 ka BP from the Lhasa Region, Tibet

This paper presents a description of the river terrace at Tangjia Village in Lhasa, Tibet. Selected types of phytolith and pollen were used as proxies to study the paleoclimate in the study area. Ancient climate and vegetation changes since 10 ka BP were examined. The results demonstrated that between 10.2 and 8.9 ka BP, the dominating phytolith was the cold type and the dominating vegetation type was grassland-forest. This indicated that the climate changed from cool-humid to cool-dry and later turned back into a cool-humid climate. Between 8.9 and 8.1 ka BP, the main types of phytoliths were tooth, dumbbell, and polyhedral. This suggests that the vegetation consisted of forest-grassland and the period＇s climate had become warmer. Between 8.1 and 6.7 ka BP, the warm index of phytolith assembelage gradually increased, whereas the spore and pollen assembelage revealed that the vegetation was forest with hardwood. This suggested that the paleoclimate was warmest in this period. The herbaceous vegetation increased gradually, indicating that the climate had become colder since 7.5 ka BP. Between 6.7 and 4.6 ka BP, cold type phytolith such as tooth and cap were found. Simultaneously, the pollen assembelage indicated that the vegetation shifted from grassland to forest and then turned back into grassland. This implies that the climate fluctuated from cold-dry to cool- humid. Between 4.6 and 1.9 ka BP, the dominate type of phytolith was cold type and its warm index was in the range 0.04-0.28, suggesting a herbaceous vegetation cover and indicating that the climate was cold. The phytolith warm index from 1.9 ka BP revealed that the climate was continuously decreasing, and most of the pollen assembelage consisted of Chenopodiaceae and Artemisia. This conclusion is in agreement with the phytolith result that indicates that the climate was becoming colder and colder.

Pilot study on binding of bovine salivary proteins to grit silicates and plant phytoliths

Mostly fed with grass in fresh or conserved form,cattle and other livestock have to cope with silicate defence bodies from plants(phytoliths)and environmental silicates(grit),which abrade tooth enamel and could additionally interact with various salivary proteins.To detect potential candidates for silicate-binding proteins,bovine whole saliva was incubated with grass-derived phytoliths and silicates.Interactions of salivary proteins with pulverized bovine dental enamel and dentine were additionally analysed.After intense washing,the powder fractions were loaded onto 1D-polyacrylamide gels,most prominent adhesive protein bands were cut out and proteins were identified by mass spectrometry within three independent replicates.All materials were mainly bound by bovine odorant-binding protein,bovine salivary protein 30×10^(3) and carbonic anhydrase VI.The phytolith/silicate fraction showed additional stronger interaction with haemoglobinβand lactoperoxidase.Conceivably,the binding of these proteins to the surfaces may contribute to biological processes occurring on them.

Pilot study on binding of bovine salivary proteins to grit silicates and plant phytoliths

Mostly fed with grass in fresh or conserved form, cattle and other livestock have to cope with silicate defence bodies from plants （phytoliths） and environmental silicates （grit）, which abrade tooth enamel and could additionally interact with various salivary proteins. To detect potential candidates for silicate-binding proteins, bovine whole saliva was incubated with grass-derived phytoliths and silicates. Interactions of salivary proteins with pulverized bovine dental enamel and dentine were additionally analysed. After intense washing, the powder fractions were loaded onto 1D-polyacrylamide gels, most prominent adhesive protein bands were cut out and proteins were identified by mass spectrometry within three independent replicates. All materials were mainly botmd by bovine odorant-binding protein, bovine salivary protein 30× 10^3 and carbonic anhydrase VI. The phytolith/silicate fraction showed additional stronger interaction with haemoglobin β and lactoperoxidase. Conceivably, the binding of these proteins to the surfaces may contribute to biological processes occurring on them.

Morphological Characteristics and Environmental Implications of Phytoliths in Topsoils from Different Vegetation Zones on Northern Slope of Changbai Mountains,China

The Changbai Mountains,located in the temperate monsoon climate zone of East Asia,is an ideal loca-tion for the research on timberline response to global changes.In this study,the topsoils were collected from different vertical vegetation zones on the northern slope of the Changbai Mountains,Northeast China in August 2009,and phytoliths in the soil samples were extracted by using wet oxidation method and identified with Motic 2.0 microscope in laboratory.The results show that phytoliths are abundant in the topsoils of the study area.The herbal phytoliths are primarily composed of elongated,tooth-shaped,point-shaped and hat-shaped phytoliths,as well as a small amount of fan-shaped and square-shaped ones.The elongated,tooth-shaped,point-shaped and hat-shaped phytoliths are representative of cold climate,while fan-shaped and square-shaped ones are representative of warm and humid climate.In the conifer broadleaved mixed forest zone,coniferous forest zone and broadleaf forest zone,there are close correlations between vegetation and woody phytoliths in the topsoils,indicating that the woody plants of a region can be reconstructed from the woody phytolith assemblages in the topsoils.Meanwhile,the topsoil phytolith assemblages can also be used to reconstruct the understory herbs effectively.The phytolith assemblages in the topsoils of the forest community and herbal community differ significantly,which can help indicate the historical location of the timberline.

Phytolith Records in Vermicular Red Earth:Implications for Bioclimatic Variationsduring Pleistocene

Phytolith analysis is employed in bioclimatic research into vermicular red earth, especially into its form, assemblage and zonation. The phytolith assemblage is divided into 10 zones. The statistics and main factor analysis of phytolith show that the obtained main factor load curves could suggest a climate change. Combined with phytolith assemblage feature, the bioclimatic variation of vermicular red earth during its formation stage, consisting of 5 arid-cold stages, 4 warm-humid stages and 2 mild stages, is reconstructed in detail. The research results indicate that phytolith records are ideal paleoclimatic signals in vermicular red earth, and that abundant information on environmental evolution can be located.

Optimization of extraction method for quantitative analysis of Si/Al in soil phytoliths

Elemental information carried by phytoliths plays a crucial indicative role in geochemical research.For instance,it serves as an indicator of the carbon pool effect in phytoliths and aids in the elucidation of silicon sources.However,early extraction methods for soil phytoliths primarily focused on obtaining their morphological and quantitative information,lacking efficient techniques for quantitative elemental analysis.In this study,we aimed to extract Si/Al information from soil phytoliths.Considering the need for complete extraction of phytolith,six extraction methods were developed and further by alkaline dissolution to determine Si/Al.Six methods were compared in terms of enrichment capacity,the weight of extracted phytoliths,and Si/Al differences.The results indicated that the addition of Ammonia-Catechol in the commonly used heavy liquid flotation method effectively improved phytolith extraction capability and the accuracy of Si/Al results.Additionally,the inclusion of an acetic acid step before alkaline dissolution further removed surface-adsorbed impurities and enhanced the analytical quality of Si/Al in phytolith.The comparison of the data in this study with other published data shows that our method is relatively robust.The improved method proposed in this study can provide a new idea for the quantitative analysis of other elements in soil phytoliths.

Sedimentary Precambrian deposits in southwestern Transbaikalia (Siberia): phytoliths content, lateral correlations and geodynamics

The Baikal Group is one of the most important stratigraphic units in the geology of East Siberia (Upper Riphean). This paper focuses on litological-facies features based on the analysis of organic remains (stromatolites, microphitolites and microfossils), in order to better precise the age and environmental deposit conditions of the Baikal Group. The results are used to draw a correlation between all the Precambrian of deposits of the South of East Siberia. The stratigraphy of the Baikal Group is then compared to the geodynamic evolution of this region, characterized by the break-up of the supercontinent Rodinia and subsequent opening of the Paleo-Asian Ocean. Stratigraphic analogs to the Baikal Group display in most areas deposits of various minerals (complex ores, phosphoresces, bauxites etc. have been dated of the same age), so the present study would also provides an economic interest.

Effects of Silicon Fertilization on Soil Chemical Properties and Phytolith Formation of Phyllostachys pubescens

Silicon is benefit to Gramineae plants in growth and resistance to various stresses.However,the effect of silicon fertilizer application on Phyllostachys pubescens is still not investigated yet.Phyllostachys pubescens Mazel ex J.Houz is one kind of Gramineae plants which distributes in a large area.In this study,a field experiment with five Si fertilizer application rates(0,125,250,375,and 500 kg ha-1)was setup in a Phyllostachys pubescens forest in China to examine the effects of Si fertilizer on bamboo Si and phytolith accumulation in fresh leaf and leaf litter.Results showed that Si application increased soil available Si content in deep layers.Si content of leaf-litter increased with the increasing level of Si fertilizer application rate,with the value ranging from 114.3 g kg^(−1)to 172.7 g kg^(−1),however,no significant difference was observed in fresh leaf,with the value ranging from 84.0 g kg^(−1)to 115.0 g kg^(−1).The phytolith contents of leaf-litter and fresh leaf were consistent with the Si contents,the phytolith content in leaf-litter of T4(500 kg ha^(−1))was 48.4%higher than the control,suggesting Phyllostachys pubescens exhibited an increasing carbon sink in phytolith when Si fertilizer applied,which is an effective way to increase long-term soil organic carbon storage in Phyllostachys pubescens forests with a suitable Si fertilization.

Climate indexes of phytoliths from Homo erectus' cave deposits in Nanjing

A study on phytoliths and their climate indexes is carried out from Homo erectus?cave deposits in HuluCave, Nanjing. Evidence shows that phytolith assemblages of the cave deposits are dominated by the cold resistant types with a lower warm index, reflecting an overall cold inclined climate. This possibly connects the cave deposits with glacial climate to a great extent, which is in accordance with thenorthern fauna revealed by fossil mammals and temperateclimate indicated by pollen assemblages. According to the distributional state of the phytoliths and their climate in-dexes on 4 profiles in the cave, it is revealed that profiles Ⅰand Ⅱ display certain cold/warm, and dry/wet fluctuations; profile Ⅲ shows a humid and cold condition with the high-est humility in the cave deposits; while profile Ⅳ indicates a possible quick accumulating process because of its stableclimate indexes except for its bottom and top.

Analysis of carbon isotope in phytoliths from C3 and C4 plants and modern soils

The analysis of carbon isotope in phytoliths from modern plants and surface soils in China shows that the values of carbon isotope are consistent with those from C3 and C4 plants, and the processes of photosynthesis of the original plants can be clearly identified by carbon isotope in phytoliths. The value of carbon isotope varied from -23.8‰ to -28‰, with the maximum distributed in the latitude zone from 34°N to 40°N in North China and East China areas, and the minimum in the Northeast China and South China regions. The values of carbon of phytoliths tend to increase from low to high and then reduce to low value again as the latitude increases. In the same latitude zone, the carbon isotope in phytoliths from grassland soil under the trees is obviously lower than that from grassland soil without any trees with the difference of 1‰ - 2‰.

Seasonal climatic variation recorded by phytolith assemblages from the Baoji loess sequence in central China over the last 150 000 a

The results from the regressive relation of phytolith assemblages from modern surface soils and climatic parameter show variations in temperature and precipitation and evolution of vegetation pattern since the last 150000 a in the Baoji loess section. The impact of changing monsoon intensity in different periods in this area is discussed.

Phytolith records of the climate change since the past 15000 years in the middle reach of the Yangtze River in China

Based on 14C dating and core sediments survey, phytolith records are employed to reconstruct paleovegetation and paleoclimate in the Jianghan Plain in the middle reach of the Yangtze River. Phytoliths identified are assigned into 21 well-described morphotypes and divided into four groups （Poaceae, fern, coniferous and broad-leaved）. The phytolith assemblages together with warmth index （Iw） are divided into 18 ecological zones, which reflect a complete vegetation history related to climate change in the middle reach of the Yangtze River during the past 15000 years. On the basis of the correlation ofphytolith records with the paleoclimatic indicators from stalagmite, peatland, North Atlantic deep-sea sediments, Loess Plateau of Central China, and Arabic Sea sediments, eight climatic phases are identified included Last Glacial Maximum （LGM） （20- 14.8 cal kaBP）, Last Deglaciation （LDG） （14.8-11.9 cal kaBP）, low-temperature phase in the Early Holocene （11.9-8 cal kaBP）, Holocene Opti- mum （8-4.9cal kaBP）, Holocene Katathermal （4.9- 1.1 cal kaBP）, Medieval Warmth Period （MWP） （1.1- 0.7 cal kaBP）, Little Ice Age （LIA） （0.7-0.15 cal kaBP）, and Modem Warming （0.15 cal kaBP-present）. Climatic events such as Bolling-Allerod warm intervals, Older Dryas, Inter-Allerod Cold Period, and Younger Dryas, and eight Holocene Bond events （B1-8） have been identified since the LDG. Our results demonstrate that the evolution of the climate in the research area has a strong link with the Indian Summer Monsoon （SW Monsoon）, Asian Summer Monsoon （SE Monsoon）, and Holocene events in North Atlantic simultaneously, which might indicate that solar variability affects the Earth surface climate system at the centennial and millennial scales.