Application of an ecosystem model for the environmental assessment of the reclamation and mitigation plans for seagrass beds in Atsumi Bay

An ecosystem model was used to evaluate the effects of reclaiming seagrass beds and creating artificial shallows with seagrass beds to mitigate the effects of the reclamation. The applied model can simulate the pelagic and benthic ecosystems including seagrass beds and tidal flats. The objectives of this study were (a) to investigate the likelihood of cultivating and maintaining seagrass beds in artificial shallows (Part 1), and (b) to understand the effects of the reclamation of seagrass beds and the creation of artificial shallows on the water quality in the estuary (Part 2). In Part 1, first, the nutrient turnover rates due to both biochemical and physical processes in the natural seagrass beds where reclamation is proposed were analyzed. Biological processes rather than physical processes were the most significant driving forces of nutrient cycles in seagrass beds. Second, the effects of filter feeding benthic fauna (suspension feeders) in the seagrass beds were analyzed. The scenario with suspension feeders resulted in higher transparency of the water column (8.7% decrease in the light attenuation coefficient) and an increase in nutrient supply (24.9% increase in NH₄-N in the water column) contributing to the high specific growth rate of seagrass. Third, the specific growth rate of seagrass on the proposed artificial shallows was measured. The value on the artificial shallows set at a depth of datum line minus 0.8 m (D.L. − 0.8 m) was approximately the same as that of the natural seagrass beds. In Part 2, first, water quality in the estuary was compared among the scenarios with/without natural seagrass beds and artificial shallows. Then, the defined values of the water purification capability of (a) artificial shallows with/without seagrass beds, and (b) natural seagrass beds per unit area were evaluated. The reclamation of the natural seagrass beds resulted in a decrease of the removal of phytoplankton and detritus from the pelagic system (i.e. resulted in a loss in the purification rate). In contrast, the creation of artificial shallows resulted in an increase of the removal of phytoplankton and detritus from the pelagic system (i.e. resulted in a gain in the purification rate). Based on an annual average, approximately twice as much phytoplankton was removed from the artificial shallows at the depth D.L. − 0.8 m, than at the depth, D.L. − 1.5 m, and the artificial shallows with seagrass beds removed pelagic DIN and DIP at a rate 120% higher than that without seagrass beds.     

Influence of giant kelp beds (Macrocystis pyrifera) on diel cycles of pCO2 and DIC in the Sub-Antarctic coastal area

The partial pressure of CO₂ (pCO₂) and dissolved inorganic carbon (DIC) were monitored in shallow coastal waters located inside and outside giant kelp beds (Macrocystis pyrifera) located in the Kerguelen Archipelago (Southern Ocean). Photosynthesis and respiration by microplankton and kelp lead to marked pCO₂ and DIC diel cycles. Daily variations of pCO₂ and DIC are significant in the spring and summer, but absent in the winter, reflecting the seasonal cycle of biological activity in the kelp beds. If the kelp beds seem to favour the onset of phytoplankton blooms, most of the primary production inside the kelp beds is due to the kelp itself. The primary production of Macrocystis kelp beds in the Sub-Antarctic high-nutrient, low-chlorophyll (HNLC) waters off the Kerguelen Archipelago is elevated and closely linked to light availability. This production is significant from October to March and reaches its climax in December at the solar radiation maximum.     

Comments on: “Underwater measurements of carbon dioxide evolution in marine plant communities: A new method” by J. Silva and R. Santos [Estuarine,Coastal and Shelf Science 78(2008) 827–830]

Silva et al. propose a new method for quantifying benthic net community production (NCP) of tidal flats under submerged condition, based on the monitoring of water pCO₂ in a transparent benthic chamber around high tide. I demonstrate here with theoretical considerations that this method is inappropriate for coastal environments, because it allows only the quantification of the change in the dissolved CO₂ which, at classical seawater pH, is only ∼10% of the change of the dissolved inorganic carbon (DIC). Total Alkalinity and/or DIC must be measured at the beginning and end of incubations in order to compute NCP in coastal environments. However, I also demonstrate that when pH is below 7, more than 95% of the DIC change occurs in the CO₂ pool. The method proposed by Silva et al. is thus valuable for freshwater environments with acidic, low alkalinity waters, where monitoring the water pCO₂ in a vial or chamber provides alone a very close approximation of the planktonic or benthic net community production.     

海洋酸化对青蛤免疫指标的影响

通过向水体中通入CO_2的方法模拟海洋酸化环境,测定青蛤(Cyclina sinensis)在酸化条件下各免疫指标的变化情况。结果显示:将青蛤置于酸化的海中(pH分别为7.4和7.7),并以自然海水做对照组(pH 8.1)后,血细胞总数随海水酸化胁迫时间的延长,表现为下降趋势,且差异显著(P0.05);海水pH降低抑制了溶菌酶的活性,但差异性不显著(P0.05);ACP活性总体呈下降趋势,对照组活性要高于酸化组,而ALP活性表现为上升趋势。酸化胁迫初期诱导SOD活性升高,后期SOD活性受到抑制,而CAT变化却截然相反;脂质过氧化产物MDA在酸化后期出现显著降低(P0.05)。     

Sedimentation within and among mangrove forests along a gradient of geomorphological settings

Coastal wetlands provide important ecological services to the coastal zone, one of which is sediment retention. In this study we investigated sediment retention across a range of geomorphological settings and across vegetation zones comprising coastal wetlands. We selected six coastal wetlands dominated by mangroves over a gradient from riverine to tidal settings in Southeast Queensland, Australia. Each site was comprised of three distinct vegetation communities distributed as parallel zones to the coast line: seaward fringe mangroves, landward scrub mangroves and saltmarsh/ cyanobacteria mat of the high intertidal zone. We measured suspended sediment retention and sedimentation rates. Additionally, in order to assess the origin of sediment transported and deposited in the mangroves, glomalin, a novel terrestrial soil carbon tracer, was used. Our results show a mean average sedimentation of 0.64 ± 0.01 mg cm⁻² spring tide⁻¹, which was variable within sites, regardless of geomorphological setting. However, geomorphological setting influenced spatial patterns of sediment deposition. Riverine mangroves had a more homogeneous distribution of sediments across the intertidal zone than tidal mangroves, where most sedimentation occurred in the fringe zone. Overall, the fringe zone retained the majority of sediment entering the coastal wetland during a tidal cycle with 0.90 ± 0.22 mg cm⁻² spring tide⁻¹, accounting for 52.5 ± 12.5% of the total sedimentation. The presence of glomalin in suspended sediments, and thus the relative importance of terrigenous sediment, was strongly influenced by geomorphological setting, with riverine mangroves receiving more glomalin in suspended solids than tidal mangroves. Glomalin was also differentially deposited within the vegetation zones at different geomorphological settings: primarily at the fringe zone of tidal mangroves and within the scrub zone of riverine mangroves. The differences we observed in the spatial distribution of sedimentation and the difference in the origin of the sediment deposited in riverine and tidal mangroves are likely to have an impact on ecological processes.     

Acidification at the Surface in the East Sea: A Coupled Climate-carbon Cycle Model Study

This modeling study investigates the impacts of increasing atmospheric CO₂ concentration on acidification in the East Sea. A historical simulation for the past three decades (1980 to 2010) was performed using the Hadley Centre Global Environmental Model (version 2), a coupled climate model with atmospheric, terrestrial and ocean cycles. As the atmospheric CO₂ concentration increased, acidification progressed in the surface waters of the marginal sea. The acidification was similar in magnitude to observations and models of acidification in the global ocean. However, in the global ocean, the acidification appears to be due to increased in-situ oceanic CO₂ uptake, whereas local processes had stronger effects in the East Sea. pH was lowered by surface warming and by the influx of water with higher dissolved inorganic carbon (DIC) from the northwestern Pacific. Due to the enhanced advection of DIC, the partial pressure of CO₂ increased faster than in the overlying air; consequently, the in-situ oceanic uptake of CO₂ decreased.     

Effects of ocean acidification and high temperatures on the bryozoan Myriapora truncata at natural CO2 vents

There are serious concerns that ocean acidification will combine with the effects of global warming to cause major shifts in marine ecosystems, but there is a lack of field data on the combined ecological effects of these changes due to the difficulty of creating large‐scale, long‐term exposures to elevated CO₂ and temperature. Here we report the first coastal transplant experiment designed to investigate the effects of naturally acidified seawater on the rates of net calcification and dissolution of the branched calcitic bryozoan Myriapora truncata (Pallas, 1766). Colonies were transplanted to normal (pH 8.1), high (mean pH 7.66, minimum value 7.33) and extremely high CO₂ conditions (mean pH 7.43, minimum value 6.83) at gas vents off Ischia Island (Tyrrhenian Sea, Italy). The net calcification rates of live colonies and the dissolution rates of dead colonies were estimated by weighing after 45 days (May–June 2008) and after 128 days (July–October) to examine the hypothesis that high CO₂ levels affect bryozoan growth and survival differently during moderate and warm water conditions. In the first observation period, seawater temperatures ranged from 19 to 24 °C; dead M. truncata colonies dissolved at high CO₂ levels (pH 7.66), whereas live specimens maintained the same net calcification rate as those growing at normal pH. In extremely high CO₂ conditions (mean pH 7.43), the live bryozoans calcified significantly less than those at normal pH. Therefore, established colonies of M. truncata seem well able to withstand the levels of ocean acidification predicted in the next 200 years, possibly because the soft tissues protect the skeleton from an external decrease in pH. However, during the second period of observation a prolonged period of high seawater temperatures (25–28 °C) halted calcification both in controls and at high CO₂, and all transplants died when high temperatures were combined with extremely high CO₂ levels. Clearly, attempts to predict the future response of organisms to ocean acidification need to consider the effects of concurrent changes such as the Mediterranean trend for increased summer temperatures in surface waters. Although M. truncata was resilient to short‐term exposure to high levels of ocean acidification at normal temperatures, our field transplants showed that its ability to calcify at higher temperatures was compromised, adding it to the growing list of species now potentially threatened by global warming.     

The effects of elevated-CO2 on physiological performance of Bryopsis plumosa

An increase in the level of atmospheric carbon dioxide (CO2) and the resultant rise in CO2 in seawater alter the inorganic carbon concentrations of seawater. This change, known as ocean acidification, ...     

Centropages typicus (Crustacea,Copepoda) reacts to volatile compounds produced by planktonic algae

Volatile organic compounds (VOCs) may play the role of infochemicals and trigger chemotaxis and ecologically relevant responses in freshwater and marine invertebrates. Aquatic grazers use these signals as chemical cues to trace the presence of their food or to detect their predators. However, detailed data are still needed to fully understand the role of these relationships in marine plankton. We investigated the ability of the copepod Centropages typicus to perceive the odour of three planktonic diatoms (Skeletonema marinoi, Pseudonitzschia delicatissima and Chaetoceros affinis) and a dinoflagellate (Prorocentrum minimum). This information is ecologically relevant for orientation, habitat selection, predator avoidance and communication. In addition, as the pH of the medium influences the perception of chemical cues in aquatic environments, we tested the effect of seawater acidification resulting from increasing levels of CO₂, and its influences on the olfactory reactions of copepods. For this reason, our tests were repeated in normal (pH 8.10) and acidified (pH 7.76) seawater in order to simulate future ocean acidification scenarios. Using replicated chemokinetic assays we demonstrated that VOCs produced by Ps. delicatissima and Pr. minimum attract copepods at normal pH, but this effect is lowered in acidified water. By contrast, the odour of S. marinoi mainly induces a reaction of repulsion, but in acidified water and at higher concentrations this toxic diatom becomes attractive for copepods. Our experiments demonstrate, for the first time, that copepods are sensitive to the volatile compounds contained in various microalgae; VOCs prompt chemokinesis according to algal species and odour concentrations. However, seawater acidification induces changes in copepods' perception of odours. These findings highlight the sensitivity of chemically mediated interactions to global changes     

Nursery fidelity,food web interactions and primary sources of nutrition of the juveniles of Solea solea and S. senegalensis in the Tagus estuary (Portugal): A stable isotope approach

Stable carbon and nitrogen isotopes were used to assess site fidelity of Solea solea and Solea senegalensis juveniles, to investigate food web interactions and to determine the dominant nutrient pathways in two nursery areas in the Tagus estuary, Portugal. Samples of water from the main sources and from the nursery areas and respective saltmarsh creeks were collected for isotope analysis, as well as sediment, benthic microalgae, saltmarsh halophytes, S. solea, S. senegalensis and its main prey, Nereis diversicolor, Scrobicularia plana and Corophium spp. While site fidelity was high in 0-group juveniles, it was lower for 1-group juveniles, possibly due to an increase in mobility and energy demands with increasing size. Analysis of the food web revealed a complex net of relations. Particulate organic matter from the freshwater sources, from each nursery's waters and saltmarsh creeks presented similar isotopic composition. Sediment isotopic composition and saltmarsh halophytes also did not differentiate the two areas. All components of the food web from the benthic microalgae upwards were isotopically different between the nursery areas. These components were always more enriched in δ¹³C and δ¹⁵N at the lower nursery area than at the nursery located upstream, appearing as if there were two parallel trophic chains with little trophic interaction between each other. A mixture of carbon and nitrogen sources is probably being incorporated into the food web. The lower nursery area is more dependent upon an isotopically enriched energy pathway, composed of marine particulate organic matter, marine benthic microalgae and detritus of the C₄ saltmarsh halophyte Spartina maritima. The two nursery areas present a different level of dependence upon the freshwater and marine energy pathways, due to hydrological features, which should be taken into account for S. solea and S. senegalensis fisheries and habitat management.     

pH variability off Goa (eastern Arabian Sea) and the response of sea urchin to ocean acidification scenarios

The increasing atmospheric CO₂ concentration in the last few decades has resulted in a decrease in oceanic pH. In this study, we assessed the natural variability of pH in coastal waters off Goa, eastern Arabian Sea. pH_T showed large variability (7.6–8.1) with low pH conditions during south-west monsoon (SWM), and the variability is found to be associated with upwelling rather than freshwater runoff. Considering that marine biota inhabiting dynamic coastal waters off Goa are exposed to such wide range of natural fluctuations of pH, an acidification experiment was carried out. We studied the impact of low pH on the local population of sea urchin Stomopneustes variolaris (Lamarck, 1816). Sea urchins were exposed for 210 days to three treatments of pH_T: 7.96, 7.76 and 7.46. Our results showed that S. variolaris at pH_T 7.96 and 7.76 were not affected, whereas the ones at pH_T 7.46 showed adverse effects after 120 days and 50% mortality by 210 days. However, even after exposure to low pH for 210 days, 50% organisms survived. Under low pH conditions (pH_T 7.46), the elemental composition of sea urchin spines exhibited deposition of excess Sr²⁺ as compared to Mg²⁺ ions. We conclude that although the sea urchins would be affected in future high CO₂ waters, at present they are not at risk even during the south-west monsoon when low pH waters reside on the shelf.     

Vulnerability of marine biodiversity to ocean acidification: A meta-analysis

The ocean captures a large part of the anthropogenic carbon dioxide emitted to the atmosphere. As a result of the increase in CO₂ partial pressure the ocean pH is lowered as compared to pre-industrial times and a further decline is expected. Ocean acidification has been proposed to pose a major threat for marine organisms, particularly shell-forming and calcifying organisms. Here we show, on the basis of meta-analysis of available experimental assessments, differences in organism responses to elevated pCO₂ and propose that marine biota may be more resistant to ocean acidification than expected. Calcification is most sensitive to ocean acidification while it is questionable if marine functional diversity is impacted significantly along the ranges of acidification predicted for the 21st century. Active biological processes and small-scale temporal and spatial variability in ocean pH may render marine biota far more resistant to ocean acidification than hitherto believed.     

Effects of low pCO2 conditions on sea urchin larval size

Ocean acidification results from an increase in the concentrations of atmospheric carbon dioxide (CO₂) impacts on marine calcifying species, which is predicted to become more pronounced in the future. By the end of this century, atmospheric pCO₂ levels will have doubled relative to the pre‐industrial levels of 280 ppm. However, the effects of pre‐industrial pCO₂ levels on marine organisms remain largely unknown. In this study, we investigated the effects of pre‐industrial pCO₂ conditions on the size of the pluteus larvae of sea urchins, which are known to be vulnerable to ocean acidification. The larval size of Hemicentrotus pulcherrimus significantly increased when reared at pre‐industrial pCO₂ level relative to the present one, and the size of Anthocidaris crassispina larvae decreased as the pCO₂ levels increased from the pre‐industrial level to the near future ones after 3 days' exposure. In this study, it is suggested that echinoid larvae responded to pre‐industrial pCO₂ levels. Ocean acidification may be affecting some sensitive marine calcifiers even at the present pCO₂ level.     

Effect of ocean acidification on coastal phytoplankton composition and accompanying organic nitrogen production

The effect of ocean acidification, caused by the increase in pCO₂ in seawater, on phytoplankton population and on related organic nitrogen production was experimentally examined by use of a natural coastal microbial population. pCO₂ and pH were controlled by aeration with air in which pCO₂ was at the current level (control), for which ambient air was used, and with air in which pCO₂ was ??800?? and ??1200?? ppm, in 500-L culture vessels. The experiment was continued for 15?days after addition of the inorganic nutrients such as nitrate, phosphate, and silicate. During most of the experimental period, a minor increase in phytoplankton biomass was noted, probably because of low irradiance, an increase in phytoplankton biomass was observed at the end of the experiment. Flow cytometric and microscopic observations revealed that this increase was because of Chrysochromulina sp. (Haptophyceae). The growth of Chrysochromulina sp. was most obvious in the control vessel, and tended to be obscured by increasing pCO₂ (decrease in pH), indicating the possibility that ocean acidification inhibits the growth of specific phytoplankton groups, for example Chrysochromulina sp. Production of particulate organic nitrogen (PON), determined by the ¹⁵N tracer method, also diminished under acidified conditions compared with that at the current level.     

The Interactive Effects of Elevated CO<Subscript>2</Subscript> and Ammonium Enrichment on the Physiological Performances of <Emphasis Type="Italic">Saccharina japonica</Emphasis> (Laminariales,Phaeophyta)

Environmental challenges such as ocean acidification and eutrophication influence the physiology of kelp species. We investigated their interactive effects on Saccharina japonica (Laminariales, Phaeophyta) under two pH conditions [Low, 7.50; High (control), 8.10] and three NH ₄ ⁺ concentrations (Low, 4; Medium, 60; High, 120 μM). The degree of variation of pH values in the culture medium and inhibition rate of photosynthetic oxygen evolution by acetazolamide were affected by pH treatments. Relative growth rates, carbon, nitrogen, and the C:N ratio in tissue samples were influenced by higher concentrations of NH ₄ ⁺ . Rates of photosynthetic oxygen evolution were enhanced under elevated CO₂ or NH ₄ ⁺ conditions, independently, but these two factors did not show an interactive effect. However, rates of NH ₄ ⁺ uptake were influenced by the interactive effect of increased CO₂ under elevated NH ₄ ⁺ treatment. Although ocean acidification and eutrophication states had an impact on physiological performance, chlorophyll fluorescence was not affected by those conditions. Our results indicated that the physiological reactions by this alga were influenced to some extent by a rise in the levels of CO₂ and NH ₄ ⁺ . Therefore, we expect that the biomass accumulation of S. japonica may well increase under future scenarios of ocean acidification and eutrophication.     

Boldness in a deep sea hermit crab to simulated tactile predator attacks is unaffected by ocean acidification

Despite rapidly growing interest in the effects of ocean acidification on marine animals, the ability of deep-sea animals to acclimate or adapt to reduced pH conditions has received little attention. Deep-sea species are generally thought to be less tolerant of environmental variation than shallow-living species because they inhabit relatively stable conditions for nearly all environmental parameters. To explore whether deep-sea hermit crabs (Pagurus tanneri) can acclimate to ocean acidification over several weeks, we compared behavioral “boldness,” measured as time taken to re-emerge from shells after a simulated predatory attack by a toy octopus, under ambient (pH ～7.6) and expected future (pH ～7.1) conditions. The boldness measure for crab behavioral responses did not differ between different pH treatments, suggesting that future deep-sea acidification would not influence anti-predatory behavior. However, we did not examine the effects of olfactory cues released by predators that may affect hermit crab behavior and could be influenced by changes in the ocean carbonate system driven by increasing CO₂ levels.     

Effects of seawater acidification by ocean CO<Subscript>2</Subscript> sequestration on bathypelagic prokaryote activities

We investigated the effects of seawater acidification induced by ocean CO₂ sequestration on bathypelagic prokaryotes. We simulated acidification conditions by bubbling high-CO₂ air or adding chemical buffer solutions to seawater samples in order to examine changes in total cell counts, heterotrophic production rate, direct viable cell count, and relative abundance of Bacteria and Archaea. Considerable suppression of prokaryotic activities was observed at pH 7.0 or lower, especially in samples enriched with organic matter. The relative abundance of Archaea increased with increasing CO₂ concentration. We found that seawater acidification can potentially alter heterotrophic activities and community structure of bathypelagic prokaryotes.     

海洋酸化对马氏珠母贝受精及早期发育的影响

自2010年7月1日至3日,在pH值为8.1、7.7和7.4条件下研究了海洋酸化对马氏珠母贝(Pinctada martensii Dunker)受精及早期发育的影响。结果显示,海洋酸化对不同pH值下马氏珠母贝的受精率无显著影响。pH8.1、pH 7.7和pH 7.4试验组幼虫的壳长、壳高的值逐渐增大,pH 8.1组幼虫的壳长、壳高的值大于同期其他两组的值,且在实验的第24、36、48小时与其他两组同期幼虫的壳长、壳高的值差异显著,这表明海洋酸化显著影响马氏珠母贝幼虫的生长。实验期间,pH 8.1试验组幼虫的存活率和畸形率没有显著变化,而pH 7.7和pH 7.4组幼虫的存活率显著低于pH 8.1组。pH 7.4组幼虫的畸形率显著高于同期pH 8.1和pH 7.7组幼虫的畸形率,表明在海洋酸化的环境中幼虫的发育受到影响。本文将为海洋酸化的相关研究提供基础数据。     

Species‐specific photosynthetic responses of symbiotic zoanthids to thermal stress and ocean acidification

Increasing sea‐surface temperatures and ocean acidification (OA) are impacting physiologic processes in a variety of marine organisms. Many sea anemones, corals and jellies in the phylum Cnidaria form endosymbiotic relationships with Symbiodinium spp. (phylum Dinoflagellata) supply the hosts with fixed carbon from photosynthesis. Much work has focused on the generally negative effects of rising temperature and OA on calcification in Symbiodinium‐coral symbioses, but has not directly measured symbiont photosynthesis in hospite or fixed carbon translocation from symbiont to host. Symbiodinium species or types vary in their environmental tolerance and photosynthetic capacity; therefore, primary production in symbiotic associations can vary with symbiont type. However, symbiont type has not been identified in a large portion of Symbiodinium?cnidarian studies. Future climate conditions and OA may favor non‐calcifying, soft‐bodied cnidarians, including zoanthids. Here we show that two zoanthid species, Palythoa sp. and Zoanthus sp., harboring different symbiont types (C1 and A4), had very different responses to increased temperature and increased partial pressure of CO₂ (pCO₂), or dissolved CO₂, and low pH. Thermal stress did not affect carbon fixation or fixed carbon translocation in the Zoanthus sp./A4 association, and high pCO₂/low pH increased carbon fixation. In contrast, both thermal stress and high pCO₂/low pH greatly inhibited carbon fixation in the Palythoa sp./C1 association. However, the combined treatment of high temperature and high pCO₂ increased carbon fixation relative to the treatment of high temperature alone. Our observations support the growing body of evidence that demonstrates that the response of symbiotic cnidarians to thermal stress and OA must be considered on a host‐specific and symbiont‐specific basis. In addition, we show that the effects of increased temperature and pCO₂ on photosynthesis may change when these two stressors are combined. Understanding how carbon fixation and translocation varies among different host?symbiont combinations is critical to predicting which Symbiodinium associations may persist in warm, acidified oceans.     

Carbon dioxide stored and acidified low oxygen bottom waters in coastal seas,Japan

Recently carbon dioxide fluxes between sea water and air have been measured in many coastal seas to clarify whether the coastal seas are source or sink of CO₂. In this study behavior of CO₂ within the water column was studied in a semi-enclosed coastal sea: the Seto Inland Sea, Japan. It was found that seasonal formation of hypoxic water mass is highly related to CO₂ dynamics in coastal seas. Bacterial remineralization of organic matter consumes dissolved oxygen (DO) and releases dissolved CO₂ in the bottom water when summertime thermal stratification develops. The CO₂ accumulates within the low DO bottom water (hypoxic water) and causes increasing of carbonic acid content which results in low pH. Concentrations of dissolved CO₂ and pH are highly correlated with DO concentration. The summertime low DO and acidification (low pH) occur in the lower layer every year. The accumulated CO₂ during the summer season is dispersed to the atmosphere at the beginning of mixing season.