Investigation of severe water problem in urban areas of a developing country: the case of Dhaka,Bangladesh

The present study evaluated water supply geochemistry in Dhaka City, Bangladesh, to provide detailed trace level (subppb) water quality data that include major ions, low dissolved oxygen (DO) and toxic trace metals for sustainable development. Dhaka Groundwater, which almost uniformly meets the World Health Organization guideline, has become the preferred source. Due to groundwater depletion and an ever-increasing need to meet water demands by city residents, Dhaka water supply and sewerage authority has initiated the treatment of river water, despite the fact that very little is known about the geochemical structure, and trace metal content in the Dhaka water supply. Major ion composition of water samples was determined, and the results used to generate Stiff diagrams. The diagrams served to visually compare water from different sources based on units of mass/volume. Hydrochemical facies analysis showed supply ground and surface waters are comprised predominately of Ca–Na–Mg–HCO₃ and Ca–Na–Mg–HCO₃–Cl types. Spatial distribution of ions, and Na/Cl and Na/SiO₂ molar ratio indicated that silicate weathering is the dominant geochemical process. Chemical data revealed that toxic Cr metal mobilization is associated with chemical hazards from the leather industry. The vulnerability of deep wells to contamination by As is governed by the geometry of induced groundwater flow paths and the geochemical conditions encountered between the shallow and deep regions of the aquifer. Quantifying total arsenic (As) and As from interlocking geochemical cycles (Fe, Mn) may assist in interpreting As dynamics in Dhaka well water. The surface source water was hypoxic to anoxic low DO associated with very high concentrations of biological oxygen demands, and electrical conductivity compared to industrial and non-industrial urban processes and standard activity guidelines. The results of this study should be applied to future research focused on the potential to improve water quality in urban and surrounding areas.     

Reconnaissance soil geochemical survey of Gibraltar

The extreme density of population of Gibraltar, situated at the southern tip of Spain, exerts considerable pressure on land use and thus future planning is of utmost importance. An initial reconnaissance soil geochemical survey of Gibraltar was based on 120 surface samples (0–15 cm) taken from a wide range of exposed, either bare soil or vegetated sites, to provide the optimum geographical distribution. The total elemental concentrations of 26 elements (Li, Na, K, Be, Mg, Ca, Sr, Ba, Al, La, Ti, V, Cr, Mo, Mn, Fe, Co, Ni, Cu, Ag, Zn, Cd, Pb, P, S, As) were determined by nitric/percholric acid digestion followed by inductively coupled plasma-atomic emission spectrometry (ICP-AES) analysis. The reconnaissance data shows that the spatial distribution of various elements depended on previous and present land use. Most elements (Ca, Cr, Mg and Mn excluded) exhibited relatively high concentrations in civilian and natural soils. Trends have been established for many elements, and concentrations exceeding guideline values have been found in certain areas of Gibraltar. This reconnaissance of Gibraltar is at present being followed by a more detailed baseline geochemical survey, which will establish the extent and magnitude of the variations in major and trace elements in soils and dusts, assess the impact of industrial, commercial and urban development on the geochemical landscape and to make recommendations concerning sustainable development.     

Enrichment of potentially toxic elements in areas underlain by black shales and slates in Korea

The Okchon black shale in Korea provides an important example of natural geological materials containing toxic elements; the Chung-Joo, Duk-Pyung, Geum-Kwan, I-Won and Chu-Bu areas are underlain by these black shales and slates of the Guryongsan Formation. This formation is part of the Okchon Group which is found in the central part of Korea. Geochemical surveys were undertaken in these five study areas in the Okchon Zone in order to examine the level of enrichment and dispersion patterns of potentially toxic elements in rocks and soils. After appropriate preparation, samples were analysed by instrumental neutron activation analysis and inductively coupled plasma atomic emission spectrometry for a range of elements. Arsenic, Cu, Mo, V, U and Zn are highly enriched in the Okchon black shales and their mean concentrations are significantly higher than those in black slates. These elements are closely associated with one another from a geochemical viewpoint and may be enriched simultaneously. Mean concentrations of As, Mo and U in soils derived from black shales occurring in the Duk-Pyung and Chu-Bu areas are higher than the permissible level suggested by Kloke (1979), and the enrichment index decreases in the order of Duk- Pyung > Chu-Bu > Chung-Joo > Geum-Kwan = I-Won areas. Uranium-bearing minerals such as uraninite and brannerite have been identified in black shales from the Chung-Joo area by electron probe micro-analysis. Uraninite grains are closely associated with monazite and pyrite with a grain size ranging from 2m to 10m whereas brannerite grains occur as a euhedral form 50m in diameter.     

Importance of background values in assessing the impact of heavy metals in river ecosystems: case study of Tisza River,Serbia

The main objective of this paper is to evaluate how a choice of different background values may affect assessing the anthropogenic heavy metal pollution in sediments from Tisza River (Serbia). The second objective of this paper is to underline significance of using geochemical background values when establishing quality criteria for sediment. Enrichment factor (EF), geoaccumulation index (I _geo), pollution load index (PLI), and potential ecological risk index (PERI) were calculated using different background values. Three geochemical (average metal concentrations in continental crust, average metal concentrations in shale, and average metal concentrations in non-contaminated core sediment samples) and two statistical methods (delineation method and principal component analyses) were used for calculating background values. It can be concluded that obtained information of pollution status can be more dependent on the use of background values than the index/factor chosen. The best option to assess the potential river sediment contamination is to compare obtained concentrations of analyzed elements with concentrations of mineralogically and texturally comparable, uncontaminated core sediment samples. Geochemical background values should be taken into account when establishing quality criteria for soils, sediments, and waters. Due to complexity of the local lithology, it is recommended that environmental monitoring and assessment include selection of an appropriate background values to gain understanding of the geochemistry and potential source of pollution in a given environment.     

Geochemical and biogeochemical features of areas of haematuria in cattle in the Eastern Carpathian Mountains of Romania

Haematuria-inducing areas in the northern and central parts of the Eastern Carpathian Mountains are characterised by the mesozoic-crystalline, neogene eruptive and transcarpathian flysch zones. The rocks have high concentrations of some metals and radioactive elements, some of them bearing polymetallic sulphides. The soils are strongly or moderately acid and oligomesobasic. They have a deficiency of soluble phosphorus and molybdenum, but up to 4.9 times the background content of metallic elements in non-affected zones. The waters have a pH around 7, a low content of salts (0.09–0.62 g L^–1), but metal levels in excess of 10 times that of the background level. Pasture vegetation has a low nutritional value, and includes up to 12% of poisonous plant species. It is characterised by low total concentrations of P, Mo, N and K, and high total concentrations of metallic trace elements, including high values of Zn and excessive values of Fe, Mn, Pb, Cr, Ni, Cd and Al. Besides clinical symptoms specific to haematuria, sick animals have abnormal blood and urinary compositions and specific morphopathological features. High concentrations of metallic elements were detected in renal calculi. The correlation of the geochemical and biogeochemical features of the haematuria-inducing areas, together with the results of the epizootological, clinical, paraclinical and morphopathological research emphasises that chronic cattle haematuria is a disease with numerous causes, but is primarily a biogeochemical problem in the area studied, making it an ecosystem disease.To whom correspondence should be addressed.     

Does the metal content in soil around a pregnant woman’s home increase the risk of low birth weight for her infant?

Low birth weight (LBW) is associated with a number of maternal environmental exposures during pregnancy. This study explored the association between soil metal concentrations around the home where the mother lived during pregnancy and the outcome of LBW. We used a retrospective cohort of 9,920 mother–child pairs who were insured by Medicaid during pregnancy and lived in ten residential areas, where we conducted soil sampling. We used a grid that overlaid the residential areas and collected soil samples at the grid intersections. The soil was analyzed for the concentration of eight metals [arsenic (As), barium (Ba), chromium (Cr), copper (Cu), lead (Pb), manganese (Mn), nickel (Ni), and mercury (Hg)], and we then used Bayesian Kriging to estimate the concentration at the actual maternal addresses, since we had the GIS coordinates of the homes. We used generalized additive modeling, because the metal concentrations had nonlinear associations with LBW, to develop the best fitting multivariable model for estimating the risk of LBW. The final model showed significant associations for female infants, maternal smoking during pregnancy, non-white mothers, Cu, and As with LBW. The As variable was nonlinear in relation to LBW, and the association between higher concentrations of As with LBW was strong (p = 0.002). We identified a statistically significant association between soil concentrations of arsenic around the home of pregnant women and an increased risk of LBW for her infant.     

Health hazard prospecting by modeling wind transfer of metal-bearing dust from mining waste dumps: application to Jebel Ressas Pb–Zn–Cd abandoned mining site (Tunisia)

This work presents a modeling approach to simulate spatial distribution of metal contamination in aerosols with evaluation of health hazard. This approach offers the advantage to be non-intrusive, less expensive than sampling and laboratory analyses. It was applied to assess the impact of metal-bearing dust from mining wastes on air quality for a nearby community and agricultural lands in Jebel Ressas (Tunisia) locality. Dust emission rates were calculated using existing parameterization adapted to the contamination source composed of mining wastes. Metal concentrations were predicted using a Gaussian model (fugitive dust model) with, as input: emission rates, dump physical parameters and meteorological data measured in situ for 30 days in summertime. Metal concentration maps were built from calculated PM10 particle concentrations. They evidence the areas where Pb and Cd concentrations exceeded WHO guidelines (0.5 and 0.005 µg/m³, respectively). Maximum concentrations of Pb and Cd in PM10 are, respectively, of 5.74 and 0.0768 µg/m³ for measured wind speed values up to 22 m/s. Preferential areas of contamination were determined in agricultural lands to the NW from the source dump where Pb and Cd exceeded guidelines up to a distance of 1,200 m. The secondary spreading directions were SW and E, toward the village. Health hazard prospecting shown that a major part of the village was exposed to contaminated dust and that daily hazard quotient (HQ) values reached locally 118 and 158, respectively, for Pb and Cd during the study period. However, HQ variations in the village are high, both temporally and geographically.     

Storm-induced transfer of particulate trace metals to the deep-sea in the Gulf of Lion (NW Mediterranean Sea)

In order to calculate budgets of particulate matter and sediment-bound contaminants leaving the continental shelf of the Gulf of Lion (GoL), settling particles were collected in March 2011 during a major storm, using sediment traps. The collecting devices were deployed in the Cap de Creus submarine canyon, which represents the main export route. Particulate matter samples were analyzed to obtain mass fluxes and contents in organic carbon, Al, Cr, Co, Ni, Cu, Zn, Cd, Pb and La, Nd and Sm. The natural or anthropogenic origin of trace metals was assessed using enrichment factors (EFs). Results are that Zn, Cu and Pb appeared to be of anthropogenic origin, whereas Ni, Co and Cr appeared to be strictly natural. The anthropogenic contribution of all elements (except Cd) was refined by acid-leaching (HCl 1 N) techniques, confirming that Zn, Cu and Pb are the elements that are the most enriched. However, although those elements are highly labile (59–77 %), they do not reflect severe enrichment (EFs <4). Most particles originate from the Rhone River. This has been confirmed by two different tracing procedures using rare earth elements ratios and concentrations of acid-leaching residual trace metals. Our results hence indicate that even in this western extremity of the GoL, storm events mainly export Rhone-derived particles via the Cap de Creus submarine canyons to the deep-sea environments. This export of material is significant as it represents about a third of the annual PTM input from the Rhone River.     

Lipids and fatty acid composition of particulate matter in the North Atlantic: importance of spatial heterogeneity,season and community structure

Lipid class profiles and total fatty acid composition of particulate matter were studied in the northeast Atlantic during the spring bloom and fall. Eddies of known physical and chemical properties were sampled at different depths. HPLC pigment data were used to characterize the phytoplankton communities. In spring, a dominance of prymnesiophytes was recorded at all depths, while in fall prochlorophytes dominated near the surface and prymnesiophytes only at deep chlorophyll maximum. Lipid classes included triglycerides, sterols, glycolipids and phospholipids. A differential relationship between phytoplankton abundance and lipid accumulation was observed: spring lipid concentrations were positively related to phytoplankton biomass, while fall particulate lipid did not show any relationship. The main feature was a northward increase in lipid concentrations unrelated to the mesoscale hydrological structures. Polar lipids dominated over neutral acyl-glycerols with phospholipids dominating over glycolipids in spring, while glycolipids dominated in fall. This resulted from different nutrient availability with a dominance of flagellates associated with mesotrophy in spring and of picophytoplankton associated with oligotrophy in fall. In terms of fatty acids, factorial correspondence analyses illustrate the influence of seasonally changing assemblages: (1) in spring, the main source of variability was the bloom with an opposition between bloom sites characterized by n-3 and n-6 PUFA, and more detrital deep samples characterized by saturated, monoenoic and branched acids; (2) fall fatty acid profiles were similar at all depths and very close to those observed for spring deep samples. Comparison of pigment and fatty acids using redundancy analysis suggested that pelagophytes were linked to saturated and branched acids. It also showed that prymnesiophytes and prochlorophytes were significantly associated with n-6 and n-3 PUFA. The spring period illustrated the complexity of these relationships with dinoflagellates and prymnesiophytes linked with n-3 PUFA, diatoms linked with palmitoleic and myristic acids, and pelagophytes linked with n-6 PUFA and higher-chain-length monoenes.     

Effects of crude oil and chemical dispersant on photosynthesis in the brain coral Diploria strigosa

An eight-hour exposure of Diploria strigosa (Dana) to a mixture of Arabian Light crude oil (19 ppm) and the chemical dispersant “Corexit 9527” (1 ppm) in a flowing seawater system reduced photosynthesis by symbiotic zooxanthellae by 85%, while either oil or dispersant alone had no effect. The greatest effect of crude oil plus dispersant occurred in the incorporation of photosynthetic products into lipids. Synthesis of was esters and triglycerides, the major storage lipids, was particularly affected. Total carbon fixation was restored within 3–5 h after treatment, and lipid synthesis was restored within 5–24 h after exposure.     

Effects of gamma-aminobutyric acid on the settlement of larvae of the black chiton Katharina tunicata

Laboratory-cultured larvae of the black chiton Katharina tunicata (Wood), collected from central California in May and June of 1979 and 1980, settled preferentially on the encrusting coralline alga Lithothamnium sp. Metamorphosis, which involves the loss of the prototrochal ciliary band, occurred within a period of 2.5 h after larvae began crawling upon the alga. In response to gamma-aminobutyric acid (GABA) at a concentration of 10^-6 M, the lecithotrophic trochophore larvae ceased swimming and settled rapidly. Developmental metamorphosis did not follow settlement in the absence of encrusting coralline alga. However, when both GABA and Lithothamnium sp. were present, larval metamorphosis occurred within 2 h of settlement. Significantly different rates of settlement exist for sibling larvae of different ages; 12 d-old larvae responded more rapily and in greater numbers to treatment with GABA than 10 d-old larvae. Differences in settling rates are attributed to the onset of metamorphic competence.     

Urban geochemistry: A study of element distributions in the soils of tallinn (Estonia)

First results are presented from the Urban geochemistry of Tallinn, a project supported by the Scientific and Environmental Affairs Division of NATO. The distribution of chemical elements in 532 samples of the topsoils from the territory of the biggest industrial centre of Estonia (pop. 500,000) is interpreted. Statistical analysis and mapping of major and trace elements at the territory of the Tallinn region and of the city were performed and background values and local anomalies of chemical elements on the territory were determined. The investigation focussed on the determination of zones with anomalously high concentrations of elements and the relationships of soil contamination with different pollution sources. The increase of the element concentrations has natural as well as anthropogenic origins. A detailed comparative analysis of the element distributions and the results of a factor analysis showed that the distribution of the major chemical elements depended mainly on the composition of the underlying sedimentary rocks.The territory of Tallinn is characterised by relatively high and widespread concentrations of Ba, Cr, Ga, Ni, Ti and Zn. Especially intensive local concentrations were determined for As, Cr, Mn, Ni, Pb, S, V and Zn, which are typical for the local pollution of the soils by industrial sources. The levels of As, Cr, Mn and V are more than three times. Pb and Zn are more than five times higher in the geochemical anomalies than for background levels. For the Tallinn region two major associations of elements connected with industrial pollutions of the soil are typical: the first association includes Ba, Cr, Mn, Ni and partly Fe and the second one includes As, Pb and Zn. For the city of Tallinn an increase of Ag, Ba, Be, La, Pb, Sn and Zn concentration in the soil was detected. Ba, Cr, Mn and Ni occur in high concentrations in the soils around Maardu. Different types of contamination sources can be identified in Tallinn and its suburbs.     

Soil organic matter and salinity affect copper bioavailability in root zone and uptake by Vicia faba L. plants

Processes that control the mobility, transformation and toxicity of metals in soil are of special importance in the root-developing zone. For this reason, there is a considerable interest in understanding trace elements (TEs) behavior in soil, emphasising the processes by which plants take them up. Increased root-zone salinity can affect plant TEs uptake and accumulation in plant tissue. Furthermore, copper (Cu) complexation by soil organic matter (SOM) is an effective mechanism of Cu retention in soils, controlling thus its bioavailability. Therefore, a greenhouse pot experiment was conducted to study the effects of soil Cu contamination in a saline environment on faba bean (Vicia faba L.) element uptake. Treatment with NaCl salinity was applied (control, 50 mM NaCl and 100 mM NaCl) on faba bean plants grown in a control and in a soil spiked with Cu (250 and 500 mg kg^?1). Low and high SOM content trial variants were studied. Cu accumulation occurred in faba bean leaf, pod and seed. Cu contamination affected plant element concentrations in leaves (Na, Ca, Mg, Mn), pod (Zn, Mn) and seed (Mn, Mo, Zn). Root-zone salinity also affected faba bean element concentrations. Furthermore, Cu contamination—salinity and salinity—SOM interactions were significant for pod Cu concentration, suggesting that Cu phytoavailability could be affected by these interactions. Future research will be focused on the mechanisms of Cu translocation in plant and adaptation aspects of abiotic stress.     

Decarboxylation of environmental L-leucine by marine red algae

The uptake and metabolism of ¹⁴C-L-leucine by Cystoclonium purpureum (Huds) Batt. (Rhodophyllidaceae) at concentrations of 0.5–50 μM were studied. Leucine was taken up from seawater at rates of 10–160 nmol·g^-1 fresh weight·h^-1. At low leucine levels ( 5 μM) the amino acid was preferably incorporated into cellular protein; amine formation was less than 5%. However at leucine concentrations of 50 μM, more than 30% of the total leucine taken up was decarboxylated to 3-methylbutylamine. This effect is expressed even more in Dumontia increassata (O. F. Müll) Lamour. (Dumontiaceae). In both species some of the amine occurred as a non-volatile derivative (bound amine) from which free amine could quantitatively be released by acid hydrolysis. Leucine and amine uptake were inhibited by CCCP. However, from cells preloaded with both leucine and amine, only free amine is released in the presence of CCCP. The results indicate that decarboxylase activity will be switched on at increased levels of amino acids in seawater which may be expected frequently in habitats of littoral algae (tidal zone). A possible function of amine formation and release within the scope of chemical defence of benthic algae is discussed.     

Trace-element speciation and partitioning in environmental geochemistry and health

Establishment of the chemical form and associations of trace elements is important in the scientific and medical fields related to environmental geochemistry and health. Fundamental understanding of trace-element behaviour, the realistic formulation of historical perspectives of trace-element contamination, an assessment of environmental transformation processes and a thorough appraisal of environment-related ill health and disease all depend on knowledge of the chemical speciation and partitioning of trace elements. These topics and the development of analytical speciation techniques and procedures are discussed with reference to trace-element studies in the Department of Forensic Medicine and Science, University of Glasgow, on lacustrine sediments and water, the atmosphere, soil and street dirt of an urban environment and human biological fluids.     

Copulation duration,but not paternity share,potentially mediates inbreeding avoidance in Drosophila montana

Studying the incidence of inbreeding avoidance is important for understanding the evolution of mating systems, especially in the context of mate choice for genetic compatibility. We investigated whether inbreeding avoidance mechanisms have evolved in the malt fly, Drosophila montana, by measuring mating latency (a measure of male attractiveness), copulation duration, days to remating, offspring production, and the proportion of offspring sired by the first (P₁) and second (P₂) male to mate in full-sibling and unrelated pairs. SNP markers were used for paternity analysis and for calculating pairwise relatedness values (genotype sharing) between mating pairs. We found 18 % inbreeding depression in egg-to-adult viability, suggesting that mating with close relatives is costly. Copulation duration was shorter between previously mated females and their brothers than with unrelated males. Based on an earlier study, shorter copulation is likely to decrease the number of inbred progeny by decreasing female remating time. However, shorter copulations did not lead to lower paternity (P₂) of full-sibling males. Progeny production of double-mated females was lower when the second male was a full-sibling as compared to an unrelated male, but we could not distinguish between inbreeding depression and lower female reproductive effort after mating with a relative. Relatedness estimates based on 34 SNPs did not detect any quantitative effect of relatedness variation on copulation duration and progeny production. We suggest that inbreeding depression has been strong enough to select for inbreeding avoidance mechanisms in our Finnish D. montana population.     

The trophic role of glycolic acid in coastal seawater. I. Heterotrophic metabolism in seawater and bacterial cultures

Glycolic acid, a known algal excretory product, represents a potentially important energy source for heterotrophic bacteria in marine waters. Measurements of heterotrophic uptake and mineralization by the natural microbes in the plankton indicate quantitative use of glycolic acid comparable to other common microbial substrates. This activity shows vertical and horizontal variations that correlate in a general way with primary productivity. Glycolic acid appears to be present at higher concentrations than other low molecular weight substrates, and it also shows a lower rate of turnover. It is mineralized (respired) to a greater extent than other recorded substrates, an average of 70% of total uptake. Two-thirds of a total of 141 colonies of bacteria cultured from seawater on marine agar proved capable oftaking up and respiring glycolic acid. These same bacteria, however, were unable to utilize glycolic acid for growth when it was the sole carbon source. Glycolate may therefore occur at higher concentrations and show erratic fluctuations in natural waters because microbial use is not tied to glycolate production but to the presence of other substrates. The fact that so many bacteria are able to metabolize it indicates a possibly important trophic role as an energy source. It is suggested that glycolic acid may be a major source of energy for active transport of other substrates by marine bacteria.     

Arsenic partitioning among particle-size fractions of mine wastes and stream sediments from cinnabar mining districts

Tailings from abandoned mercury mines represent an important pollution source by metals and metalloids. Mercury mining in Asturias (north-western Spain) has been carried out since Roman times until the 1970s. Specific and non-specific arsenic minerals are present in the paragenesis of the Hg ore deposit. As a result of intensive mining operations, waste materials contain high concentrations of As, which can be geochemically dispersed throughout surrounding areas. Arsenic accumulation, mobility and availability in soils and sediments are strongly affected by the association of As with solid phases and granular size composition. The objective of this study was to examine phase associations of As in the fine grain size subsamples of mine wastes (La Soterraña mine site) and stream sediments heavily affected by acid mine drainage (Los Rueldos mine site). An arsenic-selective sequential procedure, which categorizes As content into seven phase associations, was applied. In spite of a higher As accumulation in the finest particle-size subsamples, As fractionation did not seem to depend on grain size since similar distribution profiles were obtained for the studied granulometric fractions. The presence of As was relatively low in the most mobile forms in both sites. As was predominantly linked to short-range ordered Fe oxyhydroxides, coprecipitated with Fe and partially with Al oxyhydroxides and associated with structural material in mine waste samples. As incorporated into short-range ordered Fe oxyhydroxides was the predominant fraction at sediment samples, representing more than 80 % of total As.     

Soil and plant concentrations of cadmium and zinc in the vicinity of a smelter

Cadmium, zinc and lead concentrations in soils and plants near a smelter at Avonmouth are reported for samples collected in May 1979 and in May 1980. The total metal soil concentrations fall as distance from the smelter increases and decrease rapidly with depth for cadmium and zinc. The concentrations of these two metals are highly correlated at all depths. Near the smelter, where the carbon content and pH values of the soils are lowest, almost all the cadmium, as measured by EDTA extraction, is available. Concentration of metals in plants also decrease with distance from the smelter, with concentrations being lower in samples collected in 1980. This is possibly due to a seasonal variation in biomass, producing a dilution effect. We conclude that uptake of cadmium and that of zinc by grass species have different relationships to the carbon content of the soil.     

A light and electron microscope investigation of gametogenesis in Nicolea zostericola (Polychaeta: Terebellidae)

A light and electron microscope study was conducted on Nicolea zostericola (Oersted: Grube, 1860) to describe the processes of gametogenesis. N. zostericola is a monotelic species which spawns from 2 to 5 times over a period of 2 weeks at the end of its 1-year life span. The gonads consist of plates of cells bulging from beneath the peritoneum in the thoracic region of the body. Oogenesis has been divided into 3 phases: an ovarian phase in which gametogonia and primary oocytes are retained within the ovary, a follicular phase, in which oocytes are ovulated into the coelom surrounded by several layers of follicle cells and the vitellogenic phase, where individual oocytes rupture through the follicle cells and enter the coelomic fluid where cell growth and yolk synthesis occur. The combined ovarian and follicular phases have a duration of about 3 to 5 weeks and the vitellogenic phase about 5 to 6 months. Vitellogenesis is described on the ultrastructural level. Yolk synthesis involves the participation of the Golgi apparatus and endoplasmic reticulum of the oocyte in possible combination with nutrient absorption from the coelomic fluid. Spermatogenesis involves the rupturing of masses of spermatogonia into the coelom from the testis, surrounded by peritoneal cells. Smaller clusters of spermatocytes are later released into the coelomic fluid where mitotic and meiotic divisions eventually produce flattened plates of sperm. The mature spermatozoan possesses an elongate head, atypical for most polychaetes.