Winter Distribution of Diatom Assemblages in the East China Sea

We examined the spatial distributional relationships between diatom assemblages and water types during the winter in the East China Sea. Principal component analysis was used to identify two water types and two diatom assemblages in the study area. Coastal water types along the mainland China coastline had low temperature and salinity levels, but high nitrate levels. The shelf-mixing water type in the rest of the study area had higher temperatures and salinities and lower levels of nitrate. Diatom assemblage distribution was not spatially consistent with water type. The Kuroshio assemblage had a large standing stock, distributed along the surface of the shelf break. This assemblage is likely the result of Kuroshio surface water coming into contact with nutrient-rich water in the shelf area, triggering proliferation of certain diatom species. A background assemblage with low standing stock level persisted over the entire study area in both coastal water and the shelf-mixing water types. Our results support previous research: the background assemblage is due to poor growth conditions such as the convection of water during winter; there were no significant seasonal variations in the species composition.     

氮、磷对大亚湾大鹏澳海区浮游植物群落的影响Ⅱ种类组成

根据2005年4－5月（春季）、8月（夏季）和11月（秋季）对大亚湾大鹏澳海区表层的现场调查结合营养盐加富实验,探讨了不同季节氮（NO3－或脲氮 (urea)）和无机磷（PO43－）营养元素对浮游植物种类组成的潜在影响。大鹏澳海区浮游植物优势种有明显季节变化,秋季种类组成的平面变化最明显。春季浮游植物平均细胞密度最高但多样性指数最低。实验结果显示,春季至夏季浮游植物优势种翼根管藻模式变型Rhizosolenia alata f. genuina、丹麦细柱藻Leptocylindrus danicus和绕孢角毛藻Chaetoceros cinctus之间的演替可能受氮、磷条件变化的控制。夏季优势种菱形海线藻Thalassionema nitzschioides和威氏海链藻Thalassiosira weissflogii之间的演替可能受营养条件外的因素控制。尽管磷被认为是该海区浮游植物生长的主要限制因子,但夏季无机氮磷比值较高的实验组中未出现磷限制现象,无机氮磷比值的变化对浮游植物种类组成也没有显著影响,而秋季氮对浮游植物种类组成有较明显的潜在影响。     

Distribution of phytoplankton in the East China Sea and the southern Yellow Sea in spring in relation to environmental variables and dimethylsulfide compounds

The coastal ecosystems are highly sensitive to climate change and are usually influenced by variations in phytoplankton communities and water physiochemical factors. In the present study, the phytoplankton community, chlorophyll a（Chl a） and their relationships with environmental variables and dimethylsulfide（DMS） and dimethylsulfoniopropionate（DMSP） were investigated in spring 2017（March 24 to April 16） in the East China Sea（26.0°–33.0°N, 120.0°–128.0°E） and southern Yellow Sea（31.0°–36.0°N, 12...     

Distribution pattern of marine flagellate and its controlling factors in the central and north part of the Huanghai Sea in early summer

Abstract-A survey was carried out in the central and north part of the Huanghai Sea (34.5°～37.0°N, 120.5°～124.0°E) during June 12～27, 2000. It was found that the abundance of marineflagellate ranged from 45 to 1278 cell/ml, 479 cell/ml in average. Flagellate was more abundant in thecentral part than in the north part of Huanghai Sea, and the abundance decreased with the increasingdistance from the coast, showing a similar distribution pattern with isotherm. Vertically, high densityof flagellate was always presented in the bottom of thermocline, and formed a dense accumulation in thecentral area of the Huanghai Sea Cold Water Mass. The effects of physical and biological factors on thedistribution of marine flagellate in early summer were discussed. Water temperature (especially the ex-istence of thermocline) rather than salinity showed significant effect on the distribution pattern of marineflagellate in the Huanghai Sea in early summer. When comparing the abundance of marine flagellatewith that o     

Temporal and depth distribution of microepiphytes on Posidonia oceanica (L.) Delile leaves in a meadow off Tunisia

A survey on the epiphytic microorganisms growing on Posidonia oceanica leaves was conducted along a depth transect along the coast of Eastern Tunisia (Mahdia). Samples were collected by SCUBA diving at depths of 3, 5, 10 and 12 m in July 2008 and January 2009. A total of 58 microepiphyte taxa were identified. Multivariate analyses revealed temporal and spatial variation of the abundance of epiphytic microalgae. Water motion, light availability, temperature and motility of species seem to be responsible of temporal and bathymetric variations of epiphytic microalgae. Unlike diatoms, dinoflagellates were more abundant in deep waters, suggesting that they are more vulnerable to hydrodynamics. The significant correlation between leaf area index and abundance of epiphytic species indicates that the phenological parameters of the host plant influence the abundance of the epiphytic microorganisms. Among the epiphytic dinoflagellates, our data showed a great number of potentially toxic species (Alexandrium minitum, Amphidinium carterae, Karenia selliformis, Coolia monatis, Karlodinium veneficum, Ostreopsis siamensis, Prorocentrum concavum, Prorocentrum minimum, Prorocentrum rathymum and Prorocentrum lima). These species were more abundant in the deep station under more sheltered conditions than found at inshore shallower stations, which are subjected to high water motion.     

6种赤潮甲藻对荧光标记藻类的吞噬行为研究

选取6种在中国沿海广泛分布的赤潮甲藻米氏凯伦藻(Karenia mikimotoi)、链状亚历山大藻(Alexandrium catenella)、东海原甲藻(Prorocentrum donghaiense)、海洋原甲藻(Prorocentrum micans)、微小原甲藻(Prorocentrum minimum)和锥状斯氏藻(Scrippsiella trochoidea),采用经5-(4,6-二氯三嗪基)氨基荧光素(DTAF)标记灭活的荧光饵料藻进行投喂,观察目标甲藻是否存在吞噬行为,研究光照、营养盐条件对目标甲藻的吞噬行为的影响。结果发现,链状亚历山大藻能吞噬旋转海链藻(Thalassiosira curviseriata),东海原甲藻能摄食球等鞭金藻(Isochrysis galbana),但其摄食概率非常低,且不受光照和营养盐条件的影响。实验中,未观测到米氏凯伦藻、海洋原甲藻、微小原甲藻和锥状斯氏藻的吞噬行为。在黑暗中培养48-72h后,目标甲藻均出现不同程度的死亡,尤其是东海原甲藻和链状亚历山大藻。虽然东海原甲藻和链状亚历山大藻具吞噬行为属于混合营养生物,但光合自养是目标甲藻获取营养、维持生长最主要的方式。     

印尼混合海草床大型底栖生物丰度和多样性生态研究

海草床是海岸带最富生产力的生态系统之一,支撑着各种各样的伴生生物。热带的印度和太平洋地区被认为拥有海草植物种类多样性最高,且分布面积最广,然而,这个区域的海草床大型底栖生物我们知之甚少。为了填补认知的空白,我们在该区域开展了一项生态调查,旨在描述该区热带海草床大型底栖生物的丰度和多样性,以及确定大型底栖生物丰度、物种丰富度和群落结构是否明显存在断面内的站间变化和样地间变化。2014年5月和2015年10月我们分别在北苏拉威西省东海岸和西海岸开展野外工作,使用柱状取样器采集海草床大型底栖生物样品。所得样品共计鉴定14大类149种底栖生物,种类最为丰富的类别为多毛类（56种,占26%的总个体数）,十足类（20种,占9%的总个体数）和端足类（18种,占35%的总个体数）。东、西海岸海草床大型底栖生物表现出不同的空间分布模式。在东海岸,同一断面的大型底栖生物和端足类的丰度存在显著的站间差异;而在西海岸,大型底栖生物和多毛类的种类丰富度和丰度都表现出明显的站间变化,这可能归结于同一断面底质不均所造成。单因素ANOSIM以及MDS排序表明了北苏拉威西省东海岸和西海岸海草床大型底栖生物群落结构存在显著不同,正好对应于将海草床分成两大类型的栖息地,即西海岸的红树林-海草床-珊瑚连续体和东海岸的海草床-珊瑚连续体。与在热带海区开展的其他研究相比,本研究的大型底栖生物丰度和多样性处于中等水平。东、西海岸海草床大型底栖生物群落存在显著区别,其原因可能源于多方面,包括了沉积物模式,海草床结构和时间变化。     

Some observations on phytoplankton species composition,biomass, and productivity in Kenepuru Sound,New Zealand, 1982–83

Phytoplankton species composition, biomass, and rates of primary production were determined at two sites within Kenepuru Sound, New Zealand, in spring, summer, and autumn of 1982–83. Microflagellates and ultraplankton (< 5–10 μm) were numerically very abundant on each occasion and small gymnodinoid nanno‐planktonic (< 10–15 μm) dinoflagellates were likewise always a common component of the populations. The dinoflagellate, Prorocentrum gracile, made a substantial contribution to the total biomass in summer. The diatom community changed from mainly small chain forming species (Chaetoceros spp., Leptocylindricus spp.) in spring to small solitary centric and pennate forms (Nitzchia longissima, Coscinodiscus spp.) in summer, to a diversity of larger taxa (Coscinodiscus concinnus, Eucampia zoodiacus) in autumn. The autotrophic ciliate Mesodinium rubrum was a particularly important member of the autumn photo‐autotrophic assemblage. Both phytoplankton biomass and productivity increased from spring to autumn. In situ rates of primary production ranged from 15 to 1420 mgC m^‐2h^‐1 and chlorophyll a concentrations ranged from 6.9 to 258.5 mgChl a m^‐2. A gross primary production rate, in summer, was estimated at 0.57 gC m^‐2 d^‐1. Phytoplankton production and biomass appeared to be related to dissolved inorganic nutrient concentrations as a result of variations in the freshwater inflow. A tentative comparison between the rates of phytoplankton and cultivated mussel production is made.     

长江口及邻近海域浮游植物现存量的上下行控制分析

利用2010—2011年度3个季节的调查资料以及广义相加模型(GAM)分析,研究了长江口及邻近海域浮游植物现存量(以叶绿素a浓度表征)的上下行控制作用。调查结果显示,在叶绿素高值区,三航次营养盐含量均比整个海区平均值偏低,且春季呈现三季节最低的磷酸盐(PO4-P)和硅酸盐(SiO3-Si)浓度(平均值分别为0.48和8.96μmol/L)以及最高的氮磷比(DIN/P为43.3),为该海域春季甲藻赤潮频发提供了有利条件。夏季叶绿素高值区的硅氮比(Si/DIN)相对整个调查区较高,而春季和秋季却相近,这与夏季藻华种类主要是硅藻相一致。春、夏两季叶绿素高值区的悬浮物浓度(TSS)在时空比较上均显著低值。浮游动物高值区分布与叶绿素高值区分布虽不完全重合,但有交叉或两者相邻。GAM模型分析结果显示,各环境因子变化对长江口及邻近海域叶绿素变化的贡献可达70%以上,且主要影响因子为盐度和营养盐,而与TSS、浮游动物生物量和温度三因子的直接相关性不显著(p0.05)。受长江冲淡水的影响,盐度与DIN、PO4-P、SiO3-Si、TSS等因子间存在显著的相关关系(p0.001),说明盐度对叶绿素变化的影响可能体现了营养盐和光照条件等因子的作用。上述研究结果表明,在长江口及邻近海域,营养盐的上行效应是浮游植物现存量的主要控制作用,而光照条件和浮游动物生物量与浮游植物现存量虽然在时空分布上存在一定的联系,但非决定性控制因素。     

Heterotrophic flagellates in the littoral and sublittoral zones of the southeast part of the Pechora Sea

For the first time, data on the heterotrophic flagellate fauna in the littoral and the sublittoral zones of the southeastern part of the Pechora Sea were obtained. Sixty-six heterotrophic flagellate species were found in the study region: 48 and 42 species were found on the shelf and in the intertidal zone, respectively. The most common species were Cafeteria roenbergensis, Paraphysomonas sp., Ancyromonas sigmoides, Cafeteria minuta, Actinomonas mirabilis, and Spumella sp. The littoral fauna of the heterotrophic flagellates was more peculiar than the sublittoral fauna and equally rich. In the region studied, the community can be divided into two types: (a) the predominantly littoral community characterized by a special composition of the dominant species and by high similarity between its local varieties and (b) the heterogeneous predominantly sublittoral community characterized by a lack of a complex of particular species. The local diversity of the heterotrophic flagellate community was low (on the average, 4.5 species per one sample 1 cm³ in volume). The overall distribution pattern of the flagellates was extremely heterogeneous. The large amount of species that were encountered only once causes a great variability in the species structure from one station to another. The total number of the species monotonously increased with the growth in the number of samples and no manifested saturation of the cumulative curve was reached. This indicates the potentially greater diversity of the heterotrophic flagellate species in the region studied.     

Deep-water macrofaunal diversity in the Faroe-Shetland region (NE Atlantic): a margin subject to an unusual thermal regime

The Faroe-Shetland Channel, situated in the NE Atlantic, encompasses a number of different sediment types (habitats) as well as being subject to an unusual thermal regime. Our main objective was to assess variations in macrofaunal generic diversity and composition along two transects to gauge the relative influence of regional hydrography and local habitat heterogeneity. We found that generic richness and diversity on a West of Shetland (WoS) transect correlated most strongly with temperature range, whilst along a North of Shetland (NoS) transect, richness and diversity correlated negatively with sedimentary variables, notably total organic carbon. Macrofaunal composition at WoS is also strongly influenced by water temperature with specific genera ( e.g. Galathowenia positively associated with the temperature eigenvectors), whereas at NoS it is a combination of temperature, silt + clay fraction and total organic carbon that has an impact on composition ( e.g. Proclea and Pseudosphyrapus , showing a negative relationship with the temperature eigenvectors). Although the temperature regime exerts a strong control on regional ecology, local habitat heterogeneity remains a significant factor.     

2016年秋季热带西太平洋网采浮游植物群落结构

作者于2016年9月27日～10月25日对热带西太平洋(0°～20°N, 120°～130°E)10个站位的网采浮游植物群落结构进行了采样调查。应用Uterm?hl方法对调查海域浮游植物的物种组成、细胞丰度、优势物种以及群落多样性等相关生态特征进行了分析。希望为热带西太平洋提供一些基础的背景资料,为以后的研究奠定基础。结果表明, 鉴定出浮游植物共计4门、66属、243种(包括变种、变型), 含硅藻门(Bacillariophyta)34属、103种, 甲藻门(Pyrrophyta)28属、133种, 金藻门(Chrysophyta)2属、4种,蓝藻门(Cyanophyta)2属、3种。浮游植物细胞丰度1 965.573×10³ 细胞/m³ , 其中蓝藻的细胞丰度为1 945.169×10³ 细胞/m³ , 决定了浮游植物的分布格局, 占总细胞丰度的98.96%, 高值区分布在0°N130°E-10°N130°E的4个站位(E130-13、E130-15、E130-17、E130-19); 硅藻丰度在20°N断面N20-4站位存在高值区; 甲藻丰度在130°E断面的3个站位(E130-10、E130-13、E130-15)存在高值区。本次调查的优势种依次为铁氏束毛藻(Trichodesmium thiebaultii)、扁形原甲藻(Prorocentrum compressum)、扁豆原甲藻(Prorocentrum leniculatum)、胞内植生藻(Richelia intracellularis)、菱形海线藻(Thalassionema nitzschioides)、细弱海链藻(Thalassiosira subtilis)、具边线形圆筛藻(Coscinodiscus marginato-lineatus)、科氏角藻(Ceratium kofoidii)、鲁比膝沟藻(Gonyaulax lurbynaii)、中华半管藻(Hemiaulus sinensis)、霍氏半管藻(Hemiaulus hauckii)、小等刺硅鞭藻(Dictyocha fibula)。Shannon-Weiner多样性指数的均值为2.440,Pielou 均匀度指数的均值为0.163。相关分析结果显示浮游植物空间分布主要受PO₄-P、NH₄-N的影响,且由蓝藻的相关性决定的。聚类分析得出群落结构分为大洋群聚和近岸群聚两种类型(其中大洋群聚的站位又划分为0°～10°N纬度范围聚集和10°～20°N纬度范围聚集)。     

Spatial variability of intertidal rocky shore assemblages in the northwest coast of Portugal

The spatial variability of rocky shore assemblages of the northwest Portuguese coast was studied in a total of 12 transects, visited twice between March and August 2003. Each transect was positioned from the upper to the lower shore and four replicate samples (50 × 50 cm) were taken at each visually identified assemblage. Multivariate analysis was used to test the null hypothesis of no significant differences among assemblages located on different heights along the transect and among assemblages located at the same height on different transects. The distribution pattern of organisms along the height was not consistent across transects and significant variability could be found in assemblages located at the same height on the shore. Globally, the variability from the lower to the upper shore (vertical axis) was larger than from transect to transect (horizontal axis) but the distribution pattern along height was clearer in sheltered transects than in exposed ones. The heterogeneity in composition between the visually identified assemblages, within the same transect, was, however, the major source of variability in the study area. In the rocky intertidal northwest coast of Portugal, where tidal amplitude is broad and extremely wave-exposed sites are scarce, height above chart datum was the most important factor determining the distribution and species composition of the assemblages. However, variability along the horizontal axis was also significant and should be considered in studies of spatial patterns of distribution of organisms within this area.     

长江口海域春季浮游植物的年际变化

于2014年5月在长江口海域采集海水样品,分析了浮游植物种类组成、细胞丰度、优势种等群落结构特征。共鉴定浮游植物105种,其中硅藻30属56种,甲藻19属41种,以及隶属于金藻、蓝藻等的8个种。浮游硅藻以近岸半咸水种为主,优势种类是中肋骨条藻(Skeletonema costatum);浮游甲藻以沿岸广布种为主,典型代表是东海原甲藻(Prorocentrum donghaiense)。为了更好地了解长江口海域浮游植物的年际变化趋势,本文收集整理了该海域1986—2014年间的春季数据,分析了浮游硅藻、浮游甲藻主要群落特征的年际变化,以及浮游硅藻优势种类细胞粒径的变化趋势。结果表明:(1)浮游硅藻、浮游甲藻的种类数均呈现不同程度的上升趋势,浮游甲藻的上升幅度更为明显;(2)浮游甲藻种类数在浮游植物种类数中的占有比例呈上升趋势,而浮游硅藻所占比例呈下降趋势;(3)浮游硅藻优势种类的细胞粒径呈现小型化趋势。     

Gradient responses of epilithic diatom communities in the Baltic Sea proper

Diatom communities and assemblages are widely used as indicators of ecological change in aquatic environments and for reconstructing palaeo-environments. Good calibration data sets, directly linking changes in diatom composition to environmental factors, are needed for building reliable gradient models with high indicative value. Such models have a broad applicability because most diatom species have cosmopolitan distributions. This paper presents community analyses of brackish-water diatoms living on submerged stones in four areas of the Baltic Sea proper along the Swedish coast. The communities on the stones were composed of epilithic, epiphytic, epipelic, epipsammic and occasionally some pelagic species. Altogether, 158 quantitative samples were taken at 41 sites between 23 April and 11 May, 1990, and the data set contained 300 diatom taxa belonging to 75 genera. Species data were analysed with principal components analysis (PCA) and environmental factors were fitted passively by multiple regression analysis on the ordination results. Differences in community composition could be explained by variation in salinity (which was correlated with N:P and Si:P ratios and the occurrence of macroalgae on the stones), nutrient concentrations and variation in exposure to wave action (which was correlated to the occurrence of soft bottoms around the stones, water temperature and the occurrence of sand grains and macroalgae on the stones). Separate analyses for small taxa (cell biovolume <1000 μm³), for large taxa (cell biovolume ≥1000 μm³) and for small and large diatom taxa together showed that diatoms of different size classes respond differently to environmental variation.     

Daily Variations in Nutrient Concentrations of Seawater at 321 m Depth in Toyama Bay, Japan Sea

Nutrient concentrations (nitrate + nitrite, phosphate and silicate) in deep seawater (321 m depth) of Toyama Bay, Japan, were measured from August 1996 to July 1997 to determine the magnitude of daily variations. Significant daily variations were observed; concentrations ranged from 4.42 to 22.4 µM for nitrate + nitrite, from 0.86 to 1.98 µM for phosphate, and from 9.91 to 47.7 µM for silicate, respectively. However, there were not significant relationships between nutrient concentrations and water temperature, or between nutrients and salinity. Since temperature and salinity in the >300 m depth layer were constant through the year, the results suggest that there may be water masses with different nutrient concentrations in the deep layers (at about 320 m depth) of the bay, and a horizontal advection of these water masses may be responsible for the observed daily variations.     

赤潮异弯藻和海洋原甲藻LSU Rdna扩增及序列分析

利用引物D1R和D2C扩增并测定了赤潮异弯藻(Heterosigma akashiwo Hada)和海洋原甲藻(Prorocentrum micans Ehrenberg)的LSU rDNA D1与D2序列,并与GenBank中相关序列进行比较分析.结果表明:在种内水平,所测的H.akashiwo 6个株系之间共有5个变异位点,序列H.k与H.k-2,H.k-4均存在碱基替换;原甲藻属不同种内各株系之间的遗传距离为0.002～0.023之间,所测序列P.mi与P.micans其他株系之间均存在碱基替换.在种间水平上,原甲藻属不同种之间的遗传距离在0.045～0.139之间,大于种内遗传距离,每个种都具有特定的保守序列.根据序列间的遗传变异,可设计特异性的探针对不同株系和物种进行检测和计数.     

Plankton communities of the South Atlantic anticyclonic gyre

Data collected during cruises of the Former Soviet Union (in 1963–1989) and the British Atlantic meridional transect program (in 1995–1999) were used to analyse macroscale patterns in phyto- and zooplankton biomass, size structure, species diversity, chlorophyll a, and plankton bioluminescence in the macroscale anticyclonic gyre of the South Atlantic Ocean. The spatial pattern of bioluminescence intensity was in good agreement with that of remotely sensed (CZCS) chlorophyll a, phosphate, salinity, and copepod species diversity index distributions especially in terms of geographic inclinations of the isolines, both associated with the north-westward pattern off the South equatorial current. Among the 416 copepod species recorded in samples, 51 species were noted throughout the whole gyre. On the other hand, there were a number of species found only in one of the currents. The mesozooplankton biomass size spectra (calculated in carbon units), exhibited a fairly stable slope of the curve from the eastern periphery of the gyre to its centre. The British Atlantic meridional transect program meridional transect through the western part of the gyre showed mesozooplankton size spectra in greater detail between the equator and 50° S. Although the spectra change slowly along the transect as far as 36° S, there is a general trend toward increasing slopes from the equatorial region to the oligotrophic central gyre. The calculated phyto-to-zooplankton ratio indicated that for the tropical anticyclonic gyres, the mesozooplankton carbon biomass could be represented as the exponential function of the phytoplankton carbon.     

几株原甲藻核糖体大亚基RNA基因的部分克隆及序列分析

对7株赤潮原甲藻28S rDNA 5'端部分序列进行扩增、克隆和序列测定,并从GeneBank上获取14个原甲藻28S rDNA序列,用NJ法和ME法构建了原甲藻属的系统树,并对序列进行分析.结果表明,7株原甲藻28S rDNA扩增序列长度为950~958 bp,通过NJ法和ME法构建的系统树完全一致.大部分分离自不同海域的同种原甲藻的序列高度保守,而不同种间在序列高变区却有较大的差异.但来自南海海域的海洋原甲藻(Prorocentrum micans)与分离自其他海域的株系序列差异较大,甚至超出了有些种间的差异.由28S rDNA高变区获得的序列,有望成为浮游植物特异性分子探针设计的良好靶区域.     

Altering intertidal sediment topography: effects on biodiversity and ecosystem functioning

Sedimentary marine systems are often highly productive and perform important nutrient regeneration functions as they efficiently decompose organic material. In recent years the role of habitat effects and of species composition in ecosystem functioning has become of interest. Estuarine environments are frequently subject to considerable anthropogenic pressures whilst supporting a variety of habitats ranging from well sorted soft muds through biogenically stable sediments to highly mobile coarse sands. There is therefore considerable spatial complexity in habitat type and faunal composition. This study set out to observe the effects of altering the topographical habitat features of an estuarine mudflat on a range of porewater nutrient concentrations (NH₄⁺, NO₃⁻, NO₂⁻, PO₄³⁻ and SiO) collected from four depths (3, 5, 9, and 12 cm) and on faunal composition. Two treatments (Shelled Nets and Net Controls) were used to alter the topography from simple mud to a mussel shell crumble and were compared to un-manipulated Control areas. Sediment granulometry and organic matter content analyses alongside biological traits analysis of the fauna were also conducted.
Differences were observed in porewater nutrient concentrations between the Control and both netted treatments at 5 cm depth only; the species diversity and abundance were also different in the netted treatments compared to the Controls, although no difference between the two manipulated treatments were observed. The changes in faunal composition were attributed solely to the altered topography and the observed nutrient changes were attributed to the faunal alteration rather than the topographical manipulation.