Using diatom life-forms and ecological guilds to assess organic pollution and trophic level in rivers: a case study of rivers in south-eastern France

The European Union’s Water Framework Directive has set a target of achieving good ecological status for all aquatic environments in Europe by 2015. In order to determine the quality of aquatic environments, biological indicators such as diatoms are often used. However, biotic diatom indices can be difficult and time consuming to use because of complexity of species determination. We investigated whether the biological traits of diatoms in rivers (life-forms, size classes and ecological guilds) could be used to assess organic pollution and trophic level. We worked on a data set comprising 315 diatom species, determined at 328 river stations of south-east France and a variety of parameters. The abundances of some biological traits differed significantly between the different organic pollution and trophic levels, particularly stalked diatoms, and the motile and low-profile guilds.     

The use of periphytic diatoms as a means of assessing impacts of point source inorganic nutrient pollution in south-eastern Australia

1. Periphytic diatoms are used as indicators of water quality because their ecological tolerances or preferences to environmental variables are thought to be predictable. However, much of the present autecological information for periphytic diatoms has been derived from studies conducted in the northern hemisphere. In this present study we used periphytic diatoms to determine the impacts of inorganic nutrient pollution in a tidal river system in the temperate latitudes of south‐east Australia. In so doing, we assess the suitability of the use of the ‘northern hemisphere’ ecological tolerance/preference data for periphytic diatoms. 2. Artificial substrates were used to collect periphytic diatoms at 35 sites, which were positioned along the riverbanks and the middle of the river at various distances upstream and downstream of the sewage outfall. The sampling design took into account tidal excursions and the observed sewage plume dynamics. Periphytic diatoms were collected during the austral winter month of August and the austral spring months of September and October. We deployed the artificial substrates for 4 weeks to allow the periphytic diatoms to recruit and colonise, before identifying and enumerating the assemblages. 3. Data analysis included two approaches: multivariate visualisations of combinations of environmental and biological data to investigate shifts in species structure of the periphytic diatom assemblage and multimetric indices based on ecological tolerance/preference data. 4. We found that the spatial patterns inferred from multivariate and multimetric analyses were consistent. Temporal variation in the composition of the periphytic diatom assemblage was greater than the spatial variation along horizontal sections of the river (in any one deployment) due mainly to shifts between winter and spring species. 5. Outfall effects were most apparent in winter, possibly because subsequent deployments were swamped by growth of spring periphytic diatoms. The outfall effects included a shift towards pollutant tolerant species and a reduction in the variability of the periphytic diatom assemblage across the river. 6. We conclude that the use of periphytic diatoms and associated ecological tolerance/preference data as a means of assessing impacts of point source inorganic nutrient pollution is effective. An understanding of river and sewage flow patterns is essential to the design of appropriate monitoring programmes and to the interpretation of results, especially as periphytic diatoms are sensitive to many environmental variables.     

Epilithic Diatoms as Indicators of Water Quality in the Gravataí River, Rio Grande do Sul, Brazil

The potential use of epilithic diatoms as indicators of organic pollution was evaluated in Gravataí River, RS, (latitude 29°45′–30°12′ S; longitude 50°27′–51°12′ W). The river suffers agricultural impacts in its upper course and urban and industrial organic pollution in its lower course. Epilithic diatoms were sampled eight times from September 2000 to August 2002, at six sites. Species were identified and densities and relative abundances of populations were determined. Simultaneously, physical, chemical and microbiological variables were measured (water temperature, conductivity, turbidity, pH, dissolved oxygen, biochemical oxygen demand (BOD₅), chemical oxygen demand, ammonium, organic nitrogen, total nitrogen, ortho-phosphate, total phosphate, chloride and faecal coliforms). In order to interpret the environmental and biological variables, discriminant analysis and the TWINSPAN methods (Two-Way Indicator Species Analysis) were applied. The results indicated that the concentrations of ortho-phosphate, ammonium, total organic nitrogen, BOD₅ and faecal coliforms characterized a pollution gradient along the river, where changes in the abundance or species composition were observed. Species were classified into three groups: Group A, including species more tolerant to heavy organic pollution and eutrophication, represented by Luticola goeppertiana, L. mutica, Eolimna subminuscula, Nitzschia palea and Sellaphora pupula; Group B, comprised of tolerant and widely distributed species such as Eunotia bilunaris, Frustulia crassinervia, F. saxonica, Navicula cryptocephala, N. cryptotenella, Nitzschia palea var. tenuirostris, Surirella angusta, Pinnularia microstauron and Ulnaria ulna and Group C, with less pollution tolerant species represented by Eunotia sp. and Gomphonema parvulum.     

Freshwater fish biodiversity in the River Ganga (India): changing pattern, threats and conservation perspectives

The river Ganges is the largest river in India and the fifth longest in the world. Although, many studies on fish ecology and systematic have been conducted largely to improve fisheries but fish diversity and their distribution pattern from conservation point of view have never been adequately addressed in the Ganges. In this connection, current distribution and abundance of freshwater fishes of river Ganges was studied and assessed from April 2007 to March 2009. We documented and described 143 freshwater fish species in the all stretches of the river which is higher than what was reported earlier. Some species were observed with shift in their distribution ranges. First time, a total of 10 exotic fishes, including Pterygoplichthys anisitsi, which has never been reported from India found in the Ganges. Alterations of the hydrological pattern due to various types of hydro projects was seems to be the largest threat to fishes of Ganges. Indiscriminate and illegal fishing, pollution, water abstraction, siltation and invasion of exotic species are also threatening the fish diversity in the Ganges and as many as 29 species are listed under threatened category. The study advocates a need to identify critical fish habitats in the Ganga basin to declare them as conservation reserves to mitigate the loss of fish diversity from this mighty large river.     

Epilithic diatoms as indicators in tropical African rivers (Lake Victoria catchment)

We investigated epilithic diatoms in rivers draining to the Nyanza Bay in Lake Victoria (Kenya) with the aim of determining environmental gradients in the assemblages and testing the usefulness of diatom metrics from temperate regions. Spatial and temporal variations of assemblages were studied in 12 sites of three rivers. Kibos, Nyando, and Kisat rivers contained 224 diatom taxa collected on seven sampling occasions over 4 years. Species richness showed a marginal decrease downstream and was the lowest at sites with high conductivity and ammonia–nitrogen levels. Two-Way Indicator Species Analysis and Canonical Correspondence Analysis revealed two major groups of river sites. Conductivity, alkalinity, turbidity, dissolved oxygen, and silicate were the most important variables influencing species distribution. Ecological diatom metrics of temperate regions and the Specific Pollution sensitivity Index showed good relationships with environmental variables. Both diatom assemblages and averaged indicator values were strong in predicting sites of ecological deterioration and in differentiating an upstream site of better quality (drinking water supply of Kisumu), thereby confirming epilithic diatoms as suitable water quality indicators in equatorial rivers. The use of metrics initially designed for temperate rivers, however, appears less valuable in good quality tropical rivers because potential indicators are not considered.     

Hydrodynamic trapping in the formation of the chlorophyll a peak in turbid, very low salinity waters of estuaries

In the James River Estuary, a chlorophyll peak occurred in verylow salinity waters (0.5) during periods of low river dischargein the summer and fall. The biomass of phytoplankton, as measuredby chlorophyll concentration, was 5–10 times that formedin adjacent areas further up-and downstream. Comparisons betweenthe peak area and the 2 area for net plankton biomass, biogenicsilica, phaeopigments, POC/PON ratios and microscopic observationindicated the peak biomass was largely composed of high concentrationsof physiologically healthy freshwater diatoms. We applied equationsfor particle sinking rates to diatoms observed in the JamesRiver and compared these rates with calculations of upward verticalwater velocity. During periods of low river discharge (summerand fall) the sinking rate of diatoms in the chlorophyll peakclosely balanced the net upward water velocity, thereby selectivelytrapping diatoms in the very low salinity zone. The turbiditymaximum increased in intensity and moved downriver in winterand spring due to high river discharge. As upward water velocityincreases, phytoplankton are never able to attain a criticalbiomass and are swept downestuary. This seasonally changingplant biomass is significant to food chain and geochemical considerationsof estuaries. ¹Present address: Department of Oceanography, National FisheriesUniversity of Pusan, Korea     

Biological quality metrics: their variability and appropriate scale for assessing streams

The concept of spatial scale is at the research frontier in ecology, and although focus has been placed on trying to determine the role of spatial scale in structuring communities, there still is a further need to standardize which organism groups are to be used at which scale and under which circumstances in environmental assessment. This paper contributes to the understanding of the variability at different spatial scales (reach, stream, river basin) of metrics characterizing communities of different biological quality elements (macrophytes, fishes, macroinvertebrates and benthic diatoms) as defined by the Water Framework Directive. For this purpose, high-quality reaches from medium-sized lowland streams of Latvia, Ecoregion 15 (Baltic) were sampled using a nested hierarchical sampling design: (river basin → stream → reach). The variability of metrics within the different groups of biological quality elements confirmed that large-bodied organisms (macrophytes and fish) were less variable than small-bodied organisms (macroinvertebrates and benthic diatoms) at reach, stream and river basin scales. Single metrics of biological quality elements had the largest variation at the reach scale compared with stream and basin scales. There were no significant correlations between biodiversity indices of the different organism groups. The correlation between diversity indices (Shannon’s and Simpson’s) of the biological quality elements (macrophytes, fish, benthic macroinvertebrates and benthic diatoms) and a number of measured environmental variables varied among the different organism groups. Relationships between diversity indices and environmental factors were established for all groups of biological quality elements. Our results showed that metrics of macrophytes and fish could be used for assessing ecological quality at the river basin scale, whereas metrics of macroinvertebrates and benthic diatoms were most appropriate at a smaller scale.     

Macroinvertebrate and diatom communities as indicators for the biological assessment of river Picentino (Campania,Italy)

Pollution of rivers is an increasing problem that affects biological diversity and structure of natural ecosystem. The present study reported the results of the preliminary analysis of diatom and macroinvertebrate communities of river Picentino (Italy, Campania) in respect of the WFD/60/2000/EC. Because of the sensitive to a variety of environmental factors the two categories of organism are used as excellent indicators of changes taking place in water ecosystems, especially eutrophication. Sampling of benthic diatoms and macroinvertebrates has been carried out in five stations along the river during April–May 2014 in order to apply the ICMi and the STAR_ICMi and assess river quality. The data showed a diversification in diatomic and macroinvertebrate communities in relation to environmental stresses and level of pollution, with the disappearance of species higher sensitive to eutrophication and organic load in the upper course of the river. River water quality was found to deteriorate from the upperstream to the downstream because of the increasing of human impact and the intensive agriculture activity along the river.We conclude that the monitoring of diatomic and macroinvertebrate communities could give detailed information about the ecological status of rivers. However it is necessary to increase the achieved data by monitoring other biological communities in order to define adequated strategies to save and preserve the rivers habitat.     

Uptake and regeneration of inorganic nitrogen in coastal waters influenced by the Mississippi River spatial and seasonal variations

The Mississippi and Atchafalaya Rivers introduce large amountsof nutrients to surface waters of the northern Gulf of Mexico.This paper reports the most complete data to date on inorganicnitrogen uptake and regeneration in a broad range of coastalenvironments influenced by the river water, along with informationon nutrient concentrations and including pico-, nano-, and microplanktonspecies composition. Nitrate in surface waters is greatly reducednear the river plume, at salinities between 5 and 25 PSU, wherethe largest variance in uptake rates was observed, and was coincidentwith peaks in surface chlorophyll. Despite the depletion ofnitrate, nitrogen limitation was a rare event during the study,because of relatively high ammonium concentrations (>1 µmolNH₄⁺ I^–1 and regeneration rates. Two contrasting situationscharacterize the seasonal nitrogen dynamics in surface shelfwaters. High nitrate input during the spring caused a largebloom in which the cells were well adapted to use nitrate.Thedominant phytoplankton species were chain forming diatoms, alsoreported in sediment-trap studies in the area. Ammonium regenerationonly accounted for a small fraction of the nitrogen requirementsduring the bloom. In contrast, the low flow of river water duringsummer resulted in low nitrate concentrations in surface water.In this case phytoplankton productivity was highly reduced andmay depend greatly on ‘in sita’ ammonium regeneration.     

Epidiatomic diatoms: An insight into interaction specificity

Little is known about direct interactions between diatoms. In this study, small diatoms attaching onto larger benthic and motile diatoms were considered. This phenomenon is commonly considered as epiphytism (with diatom epiphytes) although the exact nature of this interaction has not been investigated yet. The question that was addressed here is the specificity of the interaction, simply by observing species associations in their natural environment. A comparison of this special form of epiphytism was carried out between two sites, on a Mediterranean river and one of its tributaries. The results showed both host and epiphyte selection, but with some variability in species associations. This excludes the two alternative hypotheses that (i) large diatoms are a neutral substrate for smaller diatoms and that (ii) the interactions are strictly specific between two species.     

Perspectives for the use of diatom assemblages in the water management policy of Overijssel (The Netherlands)

Diatoms have been sampled in several types of water bodies in the province of Overijssel for a period of five years. Samples from 333 sites were examined both in spring and autumn. Extensive analysis of physico-chemical variables was carried out.Data were processed by multivariate analysis for detecting correlations between species and environmental factors. Both ordination and clustering techniques were used. Statistical tests were performed to evaluate environmental relationships.The main factors in this study appear to be pH, nutrient pollution (saprobity and trophic degree) and alkalinity. Within the group of waters which is not influenced by extreme conditions of pH or pollution, there is a discrepancy between waters with low and high alkalinity.The present study confirms that diatoms are useful indicators of pollution and pH, as well documented by several authors. Diatoms are also suitable as biological indicators for alkalinity at least on a regional scale. This offers possibilities for the use of diatoms in monitoring studies, especially in waters that are still under relatively low environmental stress. It also offers opportunities for setting ecological standards, based on the characteristics of non-polluted water bodies. This aspect will be of interest to the water management policy in the province of Overijssel.     

A Multicompartment Approach - Diatoms,Macrophytes, Benthic Macroinvertebrates and Fish - To Assess the Impact of Toxic Industrial Releases on a Small French River

The River Luzou flows through a sandy substrate in the South West of France. According to the results of two assessment surveys, the Water Agency appraised that this river may not achieve the good ecological status by 2015 as required by the Water Framework Directive (2000/60/EC). This ecosystem is impacted by industrial effluents (organic matter, metals and aromatic compounds). In order to assess and characterize the impact, this study aimed to combine a set of taxonomic and non-taxonomic metrics for diatoms, macrophytes, macroinvertebrates and fish along the up- to downstream gradient of the river. Diversity metrics, biological indices, biological and ecological traits were determined for the four biological quality elements (BQE). Various quantitative metrics (biomass estimates) were also calculated for diatom communities. The results were compared to physicochemical analysis. Biological measurements were more informative than physicochemical analysis, in the context of the study. Biological responses indicated both the contamination of water and its intensity. Diversity metrics and biological indices strongly decreased with pollution for all BQE but diatoms. Convergent trait selection with pollution was observed among BQE: reproduction, colonization strategies, or trophic regime were clearly modified at impaired sites. Taxon size and relation to the substrate diverged among biological compartments. Multiple anthropogenic pollution calls for alternate assessment methods of rivers'' health. Our study exemplifies the fact that, in the case of complex contaminations, biological indicators can be more informative for environmental risk, than a wide screening of contaminants by chemical analysis alone. The combination of diverse biological compartments provided a refined diagnostic about the nature (general mode of action) and intensity of the contamination.     

Effects of in situ phosphorus enrichment on the benthos in a subalpine karst stream and implications for bioassessment in nature reserves

Park managers in nature reserves need scientifically defensible and operationally feasible ecological indicators to better manage protected areas for both nature conservation and tourism. Such needs are much more urgent in karst aquatic ecosystems where spectacular natural scenic wonders attract millions of visitors to natural areas with streams and lakes that are particularly vulnerable to nutrient pollution. To identify a set of biotic indicators of phosphorus (P) pollution in a karst stream, we conducted an in situ P enrichment experiment in a nearly pristine karst stream located at a UNESCO world heritage site in China. Our results show that both benthic algal assemblages and macroinvertebrates were sensitive to P enrichment. Changes in diatoms (e.g., Achnanthidium minutissimum, Delicata delicatula) and macroinvertebrates (e.g., collectors) were indicative of P enrichment. The color change of travertine bryophyte beds from creamy white to green was largely due to increase in filamentous green algae such as Zygnema sp., which may provide a visual cue for P enrichment and pollution. Our findings, in conjunction with further studies that directly link these indicators with human disturbance (e.g., tourist activities) throughout the park, could improve the operation of park managers in minimizing tourist-induced nutrient pollution and in operationalizing these indicators in the current environmental monitoring and assessment program.     

底栖硅藻应用于河流生态系统健康评价的研究进展

硅藻是水生态系统中最重要的初级生产者之一, 不仅为生态系统中其他消费者提供碳源, 也因分布广泛、种类多、世代时间短等特点, 通常被认为是水环境及生态系统良好的指示生物。文章介绍了河流生态系统健康及生物完整性的概念, 综述了硅藻尤其是底栖硅藻在水生态系统中的自身特点及作用; 概述了硅藻在生物监测中广泛应用的前提: 对环境因子的响应和国内外对硅藻及硅藻指数在河流生态监测中的研究进展; 介绍了广泛应用的硅藻指数及基于硅藻的生物完整性指数的原理及构建方法; 指出了硅藻及硅藻指数在生物监测及评价中可能所存在的问题, 并提出了今后我国在这个领域后期需开展的工作。     

Learning from the past: palaeohydrology and palaeoecology

Attempts to increase European biodiversity by restoring rivers and floodplains are based on inadequate data on natural systems. This is particularly the case for NW European rivers because all catchments have been impacted by agriculture and river engineering. If river restoration is to have an ecological, as opposed to `cosmetic' design basis then baseline models are required. However, this poses three questions; (a) what is the natural river‐floodplain state, (b) how can it be defined and modelled and (c) can this state be recreated today? The first two questions can only be addressed by using palaeohydrological and palaeoecological data. A second and equally vital consideration is the stability/instability of any restored system to change in external forcing factors (e.g. climate) and in this context it may not be realistic to expect baseline models to provide equilibrium solutions but instead to define process‐form domains. Over the last two decades evidence has accumulated that the natural state of lowland rivers in much of NW Europe was multi rather than single thread‐braided, anastomosing or anabranching. Until recently our knowledge of floodplain palaeoecology was generally derived from pollen diagrams, which have source‐area of problems and lack of taxonomic specificity. The precision and breadth of palaeoecological reconstruction (including richness and structure) has been greatly increased by the use of multiple palaeo‐indicators including macrofossils, diatoms and beetles. The dynamics of small to medium sized, low‐energy, predeforestation floodplains were dominated by disturbance (windthrow, beavers, etc.) and large woody debris. In order to compare the hydrogeomorphological basis of floodplain ecology, both temporally and spatially, a simple index of fluvial complexity is presented. Palaeoecological and geomorphological investigations have the potential to provide in‐depth models of the natural range of channel conditions and sensitivity to external change that can be used to provide a scientific basis for floodplain restoration. There is also the possibility that floodplain‐channel restoration may be a valuable tool in the mitigation of future geomorphological change forced by climatic instability.     

Comparative study of diatoms and macroinvertebrates as indicators of severe water pollution: Case study of the Kebena and Akaki rivers in Addis Ababa,Ethiopia

We assessed the relative performance of diatoms and macroinvertebrates to measure municipal and industrial impacts on the ecological integrity of the three major rivers flowing through Addis Ababa. Both community metric and multivariate statistical techniques were used to analyze the environmental variables and species data along the pollution gradient. This study in the Addis Ababa urban area revealed that three biologically highly stressed rivers are being impacted primarily by physical habitat degradation and both point and nonpoint pollution. The macroinvertebrate composition was liable to severe physical habitat and chemical water quality degradation. Consequently, macroinvertebrates were less diverse and not found at all at the most polluted sites with very low dissolved oxygen levels. Based on community metrics and multivariate analysis results, diatoms more reliably indicated a gradient of pollution than macroinvertebrates. However, both organism groups equally discriminated the two relatively unimpacted upstream sites from all other impacted sites. As diatoms are immobile and ubiquitous (i.e., at least a few can be found under almost any condition), they are good indicators of pollution levels among heavily impacted sites where macroinvertebrates are completely absent or less diverse. Therefore, diatoms are the powerful bioindicators for monitoring urban-impacted and seriously stressed rivers and to examine pollution gradients and impacts of specific pollution sources.     

Detecting river environmental quality through plant and macroinvertebrate bioindicators in the Aniene River (Central Italy)

From the source to the mouth of the Aniene River, a tributary of the Tiber near Rome, there is a multifactorial gradient of increasing disturbance and water pollution. We first investigated the variation of riverine and aquatic vegetation along the Aniene, analysing the floristic composition by means of transects comprising both water and shore. We then compared vascular plants and macroinvertebrates to establish which was the more reliable bioindicator of water pollution and disturbance in river habitats. The floristic dataset can be ordered according to the two axes of NMMDS. Axis I showed a high correlation with the Extended Biotic Index (EBI) (Spearman’s rho = 0.89) and Hemeroby indicator of disturbance in plants (Spearman’s rho = −0.70), and Axis II with Shannon diversity of plants (Spearman’s rho = −0.67). These results have been summarised in two GIS maps, for EBI and Hemeroby, and allow to distinguish two sectors of the Aniene: upper and low-middle, correlated with geomorphology, density, type of settlement and degree of pollution. EBI and Hemeroby indicators collect information about environmental quality along different scales.     

Comparative monitoring of diatoms, macroinvertebrates and macrophytes in the Woluwe River (Brussels, Belgium)

The river Woluwe in Brussels and Flanders (Belgium) is a small tributary of 15 km length that drains an area of 9400 ha in the Schelde river basin. The headwaters of the Woluwe are highly fragmented by diverse pond systems and are vaulted in the Brussels agglomeration. Hyporheic zones locally influence the water quality. The downstream stretch of the river receives sewage waters from households and industry. As the river Woluwe within a short distance represents a typical gradient from groundwater-fed sources in the forest towards severely polluted water, a comparative monitoring using diatoms, macroinvertebrates and macrophytes was done. The saprobic index based on diatoms, the Belgian Biotic Index (BBI) for macroinvertebrates and a macrophyte index based on the N-values of Ellenberg were used in this comparison and for estimating the correlation with the bimonthly measured chemical variables in 16 sampling stations. The diatom saprobic index and the macrophyte index were strongly correlated. Both groups showed strong correlations with phosphate, ammonium and chemical oxygen demand. The Belgian Biotic Index showed lower correlations with the nutrient variables, but was slightly better correlated to chemical oxygen demand, chloride and dissolved oxygen. None of the indices showed a correlation with nitrate. Local substrate or light conditions could interfere with the indicator system, especially for the macrophytes and occasionally for the macroinvertebrates. It was concluded that at least in this particular river system, the indices based on the primary producers were more indicative for the trophic status, whereas the BBI showed a broader relationship to the general degree of pollution. Therefore, these three indices are considered as complementary for monitoring the biological quality and the ecological status of a river system.     

Lorica Production in Choanoflagellates–A Model System for Investigating the Mechanics, Controls and Dynamics of Secretion

BIOMINERALIZATION is the process by which living organisms assemble structures from naturally occurring inorganic compounds. Mineral deposition is common and widespread amongst Protozoa and in most instances the mineralized structures provide skeletal support and protection for softer organic parts [10]. The 2 most common minerals to be deposited by Protozoa are silica and calcium carbonate. Groups of Protozoa that deposit silica, which we are concerned with here, include the diatoms, chrysophytes, choanoflagellates, Radiolar-ia, Heliozoa and testate amoebae [10]. In the majority of silica-depositing protista, silica is taken up from the medium in the form of monomelic orthosilicic acid Si(OH)₄ (soluble reactive silicate) and deposited as amorphous, polymerised biogenic silica or opal within membrane-bounded vesicles known as silica deposition vesicles (SDV). Often biogenic silica is characteristically patterned and ornamented and for most protozoan groups the morphology of silicified parts is of prime taxonomic importance. By far the most extensively studied group of silica-depositing organisms are the diatoms [1, 12, 13]. To date most of our knowledge of silica metabolism in protists has been based on investigations into this group. Diatoms require silica for the production of their frustules. Uptake and deposition of silica occurs within a closely denned portion of the cell cycle, between nuclear division and cell separation. It occupies about ± of the cell cycle and without an adequate supply of silica diatoms are unable to produce new frustule valves with the result that cell division cannot be completed. Diatoms, therefore, have an obligate requirement for silica and without this nutrient they cease to grow [11]. In contrast to diatoms a number of other silica-depositing protistan groups, such as loricate choanoflagellates and certain chrysophytes, have a facultative requirement for silica. In the past decade the ultras true ture, physiology and ecology of loricate choanoflagellates have been extensively studied by a number of different workers [7] and the significance of these studies to our understanding of the mechanisms, controls and dynamics of silica secretion is summarised and discussed here.