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	<title>state &#8211; Science</title>
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	<title>state &#8211; Science</title>
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		<title>Cellulase production by Aspergillus niger using palm kernel cake in solid state fermentation</title>
		<link>https://scienmag.com/cellulase-production-by-aspergillus-niger-using-palm-kernel-cake-in-solid-state-fermentation/</link>
		
		<dc:creator><![CDATA[Drew Townsend]]></dc:creator>
		<pubDate>Thu, 03 Sep 2026 15:37:52 +0000</pubDate>
				<category><![CDATA[Biology]]></category>
		<category><![CDATA[agricultural waste utilization]]></category>
		<category><![CDATA[Aspergillus]]></category>
		<category><![CDATA[Aspergillus niger]]></category>
		<category><![CDATA[bioethanol saccharification]]></category>
		<category><![CDATA[cake]]></category>
		<category><![CDATA[Cellulase]]></category>
		<category><![CDATA[Cellulase production]]></category>
		<category><![CDATA[cost-effective bioprocessing]]></category>
		<category><![CDATA[enzyme applications in textile and food processing]]></category>
		<category><![CDATA[enzyme manufacturing economics]]></category>
		<category><![CDATA[fermentation]]></category>
		<category><![CDATA[kernel]]></category>
		<category><![CDATA[locally sourced enzyme production]]></category>
		<category><![CDATA[niger]]></category>
		<category><![CDATA[palm]]></category>
		<category><![CDATA[palm kernel cake]]></category>
		<category><![CDATA[production]]></category>
		<category><![CDATA[regional enzyme industry development]]></category>
		<category><![CDATA[Scientific Research]]></category>
		<category><![CDATA[solid]]></category>
		<category><![CDATA[solid state fermentation]]></category>
		<category><![CDATA[state]]></category>
		<category><![CDATA[sustainable industrial enzymes]]></category>
		<guid isPermaLink="false">https://scienmag.com/?p=186363</guid>

					<description><![CDATA[None The choice of palm kernel cake as a fermentation substrate deserves closer examination in light of the broader economics of industrial enzyme manufacturing. Cellulase production at commercial scale has historically been dominated by a small number of multinational suppliers,]]></description>
										<content:encoded><![CDATA[<p>None<br />
The choice of palm kernel cake as a fermentation substrate deserves closer examination in light of the broader economics of industrial enzyme manufacturing. Cellulase production at commercial scale has historically been dominated by a small number of multinational suppliers, and the cost of the carbon source can account for a substantial fraction of total production expenses. When enzyme producers in importing countries must ship finished enzyme preparations or refined substrates across long distances, the final price paid by downstream users in West Africa rises considerably. By demonstrating that a locally abundant by-product can support meaningful cellulase titers, the study addresses a structural problem rather than merely a technical one. Palm kernel cake is generated in large volumes wherever palm kernel oil is pressed, and in the Ashanti Region of Ghana it is often available at little more than the cost of transport. Converting this material into enzyme-rich fermented biomass would allow small and medium enterprises to produce crude enzyme preparations for local applications such as textile processing, food clarification, or the saccharification of agricultural residues for bioethanol, without the burden of import duties and cold-chain logistics.</p>
<p>The temporal profile reported in the study also illustrates a well-known feature of fungal physiology in solid-state culture. Enzyme secretion by filamentous fungi typically follows a pattern in which activity rises during the early exponential growth phase, peaks as the fungus encounters nutrient limitation or accumulates metabolic by-products, and then declines as proteolysis, substrate depletion, or product inhibition erode the accumulated enzyme pool. The observed rise from 4.68 units per milliliter on day two to 11.96 units per milliliter on day four, followed by a gradual decline to roughly 8.5 to 8.9 units per milliliter by days eight and ten, is consistent with this classical pattern. The rapid early increase suggests that the fungus germinated quickly on the moistened cake and that the cellulose fraction of the substrate was sufficiently accessible to induce cellulase synthesis. Induction of cellulase genes in Aspergillus species is generally triggered by the presence of cellulose or soluble cellulose derivatives, while high concentrations of glucose repress expression through carbon catabolite repression. The fact that activity climbed steeply between days two and four implies that the readily available sugars in the palm kernel cake were consumed early, relieving repression and allowing the cellulolytic system to be fully expressed.</p>
<p>The decline in activity after the day-four peak can be interpreted through several non-exclusive mechanisms. Proteases secreted by the fungus during later growth stages can degrade cellulases, particularly in substrates containing modest nitrogen reserves. The physical structure of the substrate also changes over time; as the fungus consumes the more digestible polysaccharide fractions, the remaining material becomes increasingly recalcitrant, reducing the stimulus for continued enzyme production. Additionally, the accumulation of soluble hydrolysis products can exert feedback inhibition on the enzymes themselves, and the crude extract measured at each time point reflects a dynamic balance between ongoing secretion and in situ degradation. The stabilization observed between days eight and ten may indicate that the culture had entered a quiescent state in which residual enzyme activity persisted without further net production. For process design, this plateau is informative because it suggests a window of several days during which harvest would yield comparable activity, providing operational flexibility even though the statistical analysis identified day four as the optimum.</p>
<p>The assay methodology used in the study merits some elaboration for readers less familiar with cellulase measurement. The dinitrosalicylic acid method detects reducing sugars released from a substrate, and when carboxymethylcellulose is used as the substrate, the measured activity corresponds primarily to endoglucanase activity. Endoglucanases cleave internal glycosidic bonds within amorphous regions of cellulose chains, generating new chain ends and reducing polymer length. This is only one component of the complete cellulolytic system, which also includes exoglucanases that processively release cellobiose from chain ends and beta-glucosidases that cleave cellobiose into glucose. A full characterization of the secreted enzyme cocktail would require additional assays, such as filter paper activity for total cellulase capacity or p-nitrophenyl-beta-D-glucopyranoside assays for beta-glucosidase. Because palm kernel cake contains substantial hemicellulose in addition to cellulose, the fungus likely secreted xylanases and other accessory enzymes as well, and these could be valuable co-products in their own right. The single-enzyme measurement reported therefore represents a conservative lower bound on the total hydrolytic value of the crude extract.</p>
<p>The normalization of activity to dry substrate weight, expressed as units per gram of dry substrate, is an important methodological detail that facilitates comparison across the solid-state fermentation literature. Reported cellulase yields on lignocellulosic residues vary widely depending on substrate composition, particle size, moisture regime, inoculum density, and fungal strain, and expressing results per gram of dry substrate rather than per milliliter of extract removes one major source of ambiguity. The extraction procedure used, involving shaking with distilled water followed by centrifugation, is a standard approach for recovering extracellular enzymes from fermented solids, though extraction efficiency is rarely complete and can itself depend on incubation time as the enzyme distribution between solid and liquid phases shifts. Readers comparing these results with other studies should therefore attend to whether activities were measured in crude extracts or in culture filtrates, and whether the assay temperature and pH matched those used here.</p>
<p>The moisture content of eighty percent deserves comment because water activity is among the most influential parameters in solid-state fermentation. Filamentous fungi tolerate lower water activities than bacteria, which is one reason solid-state culture favors fungal enzyme production and reduces bacterial contamination risk. Too little water limits swelling of the substrate, diffusion of nutrients, and mass transfer of secreted enzymes; too much water fills the interparticle spaces, reduces oxygen availability, and can effectively convert the process toward submerged conditions with their attendant disadvantages. The level chosen in this study sits within the range commonly reported for fungal solid-state fermentation of lignocellulosic substrates, but it was held constant, so the interaction between moisture and incubation time remains unexplored. Similarly, the ambient laboratory temperature, which in Kumasi typically falls in the mid-twenties to low thirties Celsius, was not actively controlled, meaning that the reported kinetics reflect a realistic but variable thermal environment. Future work that systematically varies moisture, temperature, particle size, and inoculum density, ideally through a factorial or response-surface design, would be needed to identify the true optimum and to quantify interactions among these parameters.</p>
<p>The absence of an uninoculated control, which the authors acknowledge, is worth considering from the standpoint of experimental interpretation. Because the substrate was autoclaved before inoculation, the contribution of native microbial communities to the measured activity is likely minimal, and the steep rise in activity coinciding with fungal growth supports the attribution of cellulase production to Aspergillus niger. Nevertheless, palm kernel cake may contain residual enzymes or heat-stable reducing sugars that could contribute to background reducing sugar release in the assay, and a substrate-only control would have allowed this background to be subtracted. Similarly, a heat-inactivated or killed-mycelium control would help distinguish enzyme activity from abiotic sugar release. These considerations do not undermine the central finding, given the clear temporal dynamics, but they define the boundaries within which the quantitative values should be interpreted.</p>
<p>From an applied perspective, the crude enzyme produced on palm kernel cake would be most immediately useful in applications that tolerate impurities and variable composition. In textile bioprocessing, bio-polishing of cotton fabrics, and in the softening of denim, crude cellulase preparations are routinely used and substrate-derived impurities are of limited concern. In the saccharification of agricultural residues for bioethanol, the enzyme cocktail would act on the same class of substrates on which it was induced, potentially providing good activity against local residues such as cassava peels, cocoa pod husk fiber, or rice husk. The co-secretion of xylanases by the fungus would be advantageous in these contexts because hemicellulose often shields cellulose fibers and its removal improves overall hydrolysis efficiency. For higher-value applications such as pharmaceutical or food-grade enzymes, additional purification and quality assurance would be required, but the fermentation step demonstrated here would remain the foundation of the process.</p>
<p>The circular economy framing of the work connects it to a wider agenda of agro-industrial waste valorization across West Africa. Palm oil processing generates several distinct residue streams, including empty fruit bunches, palm oil mill effluent, palm kernel shells, and palm kernel cake, each with different composition and potential uses. Palm kernel cake is already used to a limited extent as animal feed because of its residual protein and fat content, but its high fiber fraction limits its digestibility for monogastric animals. Enzymatic treatment or solid-state fermentation of the cake could serve a dual purpose: producing cellulase as a harvestable product while simultaneously upgrading the residual solid as a feed ingredient with improved fiber digestibility. Such integrated biorefinery concepts, in which a single substrate yields sequential or co-located products, are increasingly viewed as the most economically viable route for small-scale biotechnology in developing regions, because no single low-value product carries the entire process cost.</p>
<p>Finally, the study&#8217;s identification of day four as the optimal harvest point provides a concrete anchor for subsequent optimization and scale-up efforts. In industrial practice, the productivity of a fermentation process, expressed as units produced per liter per day, often matters more than the peak titer itself, and a short four-day cycle with high activity compares favorably with longer fermentations that achieve only modestly higher peaks. The single-factor design used here is an appropriate first step, establishing the temporal baseline against which the effects of other variables can be measured. As multifactorial studies build on this foundation, and as strain improvement and medium supplementation are explored, the combination of a GRAS-status producer, a locally sourced substrate, and a defined production window offers a credible starting point for developing cellulase production capacity within Ghana and, by extension, a template applicable to other palm-processing regions facing similar waste management and enzyme import challenges.</p>
<p><strong>Subject of Research:</strong> Cellulase production by Aspergillus niger using palm kernel cake in solid state fermentation</p>
<p><strong>Article Title:</strong> Cellulase production by Aspergillus niger using palm kernel cake in solid state fermentation</p>
<p><strong>Article References:</strong> Sam Barko, P., Cofie, H., Danquah, B., Bentil, J. A., &amp; Ofosu, M. (2026). Cellulase production by Aspergillus niger using palm kernel cake in solid state fermentation. <em>Discover Biotechnology, 3</em>(1), Article 10. <a href="https://doi.org/10.1007/s44340-026-00056-z" rel="noopener noreferrer">https://doi.org/10.1007/s44340-026-00056-z</a></p>
<p><strong>Image Credits:</strong> AI Generated</p>
<p><strong>DOI:</strong> <a href="https://doi.org/10.1007/s44340-026-00056-z" rel="noopener noreferrer">10.1007/s44340-026-00056-z</a></p>
<p><strong>Keywords:</strong> Cellulase, production, Aspergillus, niger, palm, kernel, cake, solid, state, fermentation, scientific research</p>
]]></content:encoded>
					
		
		
		<post-id xmlns="com-wordpress:feed-additions:1">186363</post-id>	</item>
		<item>
		<title>India’s Coastal Waters Reveal Emerging Ocean Acidification Hotspots</title>
		<link>https://scienmag.com/indias-coastal-waters-reveal-emerging-ocean-acidification-hotspots/</link>
		
		<dc:creator><![CDATA[Violet Maxwell]]></dc:creator>
		<pubDate>Fri, 28 Aug 2026 23:45:53 +0000</pubDate>
				<category><![CDATA[Earth Science]]></category>
		<category><![CDATA[Aragonite]]></category>
		<category><![CDATA[aragonite saturation]]></category>
		<category><![CDATA[carbonate chemistry]]></category>
		<category><![CDATA[coastal marine ecosystems]]></category>
		<category><![CDATA[coral reef vulnerability]]></category>
		<category><![CDATA[coral reefs]]></category>
		<category><![CDATA[early warning signals for ocean health]]></category>
		<category><![CDATA[fisheries impacts]]></category>
		<category><![CDATA[Gulf of Mannar]]></category>
		<category><![CDATA[Gulf of Mannar ecological study]]></category>
		<category><![CDATA[Indian Ocean]]></category>
		<category><![CDATA[indicates]]></category>
		<category><![CDATA[local versus global ocean acidification]]></category>
		<category><![CDATA[Marine Conservation Planning]]></category>
		<category><![CDATA[ocean acidification]]></category>
		<category><![CDATA[Palk Bay]]></category>
		<category><![CDATA[Palk Bay carbonate chemistry]]></category>
		<category><![CDATA[saturation]]></category>
		<category><![CDATA[seagrass meadow health]]></category>
		<category><![CDATA[seagrass meadows]]></category>
		<category><![CDATA[seasonal variability in ocean chemistry]]></category>
		<category><![CDATA[shellfish calcification]]></category>
		<category><![CDATA[state]]></category>
		<guid isPermaLink="false">https://scienmag.com/?p=184153</guid>

					<description><![CDATA[A seasonal survey found lower aragonite saturation and pH in India’s Gulf of Mannar than in Palk Bay, identifying the former as more vulnerable to ocean acidification.]]></description>
										<content:encoded><![CDATA[<p>Two ecologically important coastal waters along southeastern India are showing markedly different chemical conditions that could shape the future of coral reefs, seagrass meadows, shellfish and fisheries. A study of the Gulf of Mannar and nearby Palk Bay found that the Gulf of Mannar had lower average pH and lower aragonite saturation state, a combination that signals greater vulnerability to ocean acidification. The research examined seawater collected during four seasons from 24 stations across the two semi-enclosed marine systems in 2023 and 2024. Although neither region had reached the chemical point at which aragonite dissolves outright, the Gulf of Mannar repeatedly approached less favorable conditions for organisms that build shells and skeletons from calcium carbonate. Palk Bay, by comparison, generally retained a larger chemical margin for calcification, although its carbonate chemistry varied more strongly through the year. The findings place local ocean chemistry at the center of conservation planning for two habitats whose ecological value extends well beyond their shorelines. They also demonstrate why broad global averages can miss the early warning signals emerging in dynamic coastal seas.</p>
<p>Ocean acidification begins when carbon dioxide from the atmosphere dissolves into seawater. The gas reacts with water to form carbonic acid, which releases hydrogen ions and shifts the balance among dissolved carbon species. As hydrogen-ion concentrations rise, pH falls and carbonate ions become less available. Those ions are essential ingredients for organisms that construct aragonite or calcite, two crystalline forms of calcium carbonate. The aragonite saturation state, written as Ω<sub>arag</sub>, summarizes how favorable the water is for forming aragonite. Values above one indicate supersaturation, while values below one indicate undersaturation, when dissolution can become thermodynamically favored. Yet remaining above one does not mean that all calcifying organisms are unaffected. Growth and calcification can become more difficult well before waters become undersaturated, especially when acidification occurs alongside heat, low oxygen, pollution or nutrient enrichment. Because Ω<sub>arag</sub> responds to pH, carbonate-ion concentration, dissolved carbon dioxide and the ocean’s buffering capacity, it can reveal ecological stress that a pH measurement alone may not fully capture.</p>
<p>The study area contains a dense mosaic of habitats. The Gulf of Mannar stretches between Tuticorin and Mandapam and includes 21 islands surrounded by coral reefs, mangroves and seagrass. Its waters support an extraordinary variety of marine life, including fishes, mollusks and reef-associated invertebrates. Palk Bay is a shallow, semi-enclosed basin connected to the Bay of Bengal and strongly influenced by river-borne sediments and freshwater. Both systems are shaped by the seasonal monsoon, but their depth, circulation, sediment transport and biological communities differ. Those differences can alter how quickly carbon dioxide accumulates, how efficiently waters mix and how much carbonate remains available. The researchers selected stations near coral reefs, seagrass meadows and mangrove ecosystems to capture this environmental range. They collected subsurface samples at depths of roughly 0.5 to 1 meter during the Northeast Monsoon, Post-Monsoon, Summer and Southwest Monsoon. Each station was sampled in triplicate, allowing the team to assess both regional patterns and the precision of its measurements.</p>
<p>The contrast between the regions was clearest in their average carbonate conditions. Palk Bay recorded a mean pH of 8.33 plus or minus 0.06, compared with 8.08 plus or minus 0.02 in the Gulf of Mannar. Its mean Ω<sub>arag</sub> reached 3.22 plus or minus 0.57, while the Gulf of Mannar averaged 2.82 plus or minus 0.20. These values remain above the saturation threshold, but the lower Gulf of Mannar average indicates less favorable conditions for calcium-carbonate production. The researchers identified particularly low Ω<sub>arag</sub> values, below three, during the Post-Monsoon season at the Kurusadai and Vedalai stations in the Gulf of Mannar. Palk Bay remained above three during the same season. The distinction is not a forecast of immediate reef collapse, nor does it establish a biological threshold for every species. Instead, it identifies a chemical gradient: organisms in the Gulf of Mannar may have less energy available for skeletal growth and less resilience when acidification is combined with warming or other disturbances.</p>
<p>Seasonal changes were driven by a shifting mixture of physical and biological processes. During the Northeast Monsoon, average surface temperatures were about 29.7 degrees Celsius in Palk Bay and 29.0 degrees in the Gulf of Mannar. By Summer, both regions approached 31.8 degrees. Salinity also rose during Summer, reaching an average of 35.83 practical salinity units in Palk Bay and 34.43 in the Gulf of Mannar, compared with lower values during the Northeast Monsoon. Monsoon winds alter circulation, freshwater delivery, sediment movement and vertical mixing. The Southwest Monsoon can transport upwelled, carbon-dioxide-rich water toward the Gulf of Mannar, while the Northeast Monsoon can carry lower-salinity water from the Bay of Bengal. Freshwater and nutrients can modify alkalinity and biological productivity, while respiration and the decomposition of organic material can add carbon dioxide to coastal waters. Photosynthesis can temporarily remove carbon dioxide near seagrass and algal communities, raising pH and carbonate availability. These competing influences help explain why the same coastline can experience sharp seasonal swings rather than a uniform, steadily declining signal.</p>
<p>Measurements of the carbonate system supported that interpretation. In Palk Bay, mean seawater partial pressure of carbon dioxide ranged from about 141 to 241 microatmospheres across seasons; in the Gulf of Mannar, the range was approximately 188 to 225 microatmospheres. The highest Palk Bay average occurred during the Northeast Monsoon, when freshwater inputs and mixing may have reshaped the local carbon balance. Carbonate-ion concentrations were generally higher in Palk Bay than in the Gulf of Mannar, while the Revelle factor, a measure related to the ocean’s resistance to absorbing additional carbon dioxide, ranged from 6.85 to 7.83 in Palk Bay and 7.39 to 7.74 in the Gulf of Mannar. A higher Revelle factor means that a given increase in dissolved inorganic carbon can produce a comparatively larger rise in seawater carbon dioxide. The researchers calculated carbonate variables with the CO<sub>2</sub>SYS program using laboratory pH, temperature, salinity, total alkalinity, phosphate and silicate measurements. This approach allowed them to estimate pCO<sub>2</sub>, carbonate ions, calcite saturation, aragonite saturation and buffering-related properties from a consistent set of chemical observations.</p>
<p>Statistical analyses pointed to carbonate chemistry, rather than any single physical measurement, as the principal control on Ω<sub>arag</sub>. A two-way analysis of variance found significant effects of both season and region, as well as a significant interaction between them, meaning that the magnitude of seasonal variability differed between Palk Bay and the Gulf of Mannar. Pearson correlations showed a moderate positive relationship between Ω<sub>arag</sub> and pH, with a correlation coefficient of 0.672, and a much stronger relationship with carbonate-ion concentration, with a coefficient of 0.959. Ω<sub>arag</sub> was negatively related to pCO<sub>2</sub> and the Revelle factor. The team also used structural equation modelling to examine direct and indirect links among 15 environmental variables. In that model, pH and carbonate ions exerted strong positive influences on aragonite saturation, while pCO<sub>2</sub> exerted a negative influence. Temperature, salinity and nutrient concentrations played smaller or indirect roles. The analysis reinforces a basic chemical principle: when excess carbon dioxide shifts carbonate ions toward bicarbonate, calcifying organisms face a reduced supply of the building blocks needed for aragonite.</p>
<p>The ecological consequences could reach across the food web and into coastal economies. Coral reefs create three-dimensional habitat for fish and invertebrates, shelter young organisms and support fisheries. Mollusks and echinoderms also depend on calcium-carbonate structures, and previous experimental work has shown that tropical sea urchins can be sensitive to carbon-dioxide-driven changes in calcification and physiology. A weaker balance between reef construction and erosion could gradually reduce habitat complexity, even if seawater remains technically supersaturated. The study suggests that Palk Bay’s extensive seagrass meadows may help moderate local conditions by taking up carbon dioxide during photosynthesis and storing carbon in biomass and sediments. The Gulf of Mannar has less seagrass coverage in some areas, and habitat degradation and sedimentation may reduce this potential buffer. Its deeper waters and exposure to monsoon-linked upwelling may further increase the delivery of carbon-dioxide-rich water. The researchers therefore describe the Gulf of Mannar as more vulnerable than Palk Bay, while emphasizing that both systems require continued observation. Their recommended next step is long-term monitoring that combines Ω<sub>arag</sub>, pH, pCO<sub>2</sub>, alkalinity, nutrients, temperature, oxygen and biological surveys, providing managers with an early-warning system for changing coastal conditions.</p>
<p>These results should be interpreted as a baseline rather than as a long-term trend. The investigation was a pilot assessment covering 24 locations during 2023–2024, so repeated observations over many years will be needed to distinguish persistent acidification from normal coastal variability. That distinction matters particularly in semi-enclosed waters, where river discharge, sediment movement, monsoon circulation and biological carbon cycling can change carbonate chemistry over short distances and time periods. A single regional average may therefore conceal conditions experienced by organisms living near an island reef, seagrass meadow or sediment-influenced shoreline.</p>
<p>Aragonite saturation is also best viewed alongside measurements of total alkalinity, dissolved inorganic carbon, oxygen, nutrients and temperature. Together, these variables can help identify whether low saturation reflects atmospheric carbon dioxide uptake, respiration and organic-matter decomposition, freshwater dilution, upwelling or changes in alkalinity. Biological surveys are equally important because species differ in their sensitivity and capacity to acclimate. Tracking coral growth, mollusk recruitment, echinoderm abundance and seagrass condition with carbonate chemistry would link chemical exposure to ecosystem response. Such integrated observations could help separate areas that are naturally variable from emerging hotspots where local stressors amplify the broader influence of rising carbon dioxide.</p>
<p><strong>Subject of Research:</strong> Seasonal ocean acidification vulnerability in the Gulf of Mannar and Palk Bay</p>
<p><strong>Article Title:</strong> Aragonite saturation state indicates emerging ocean acidification hotspots in the Gulf of Mannar and Palk Bay along the Southeast Coast of India</p>
<p><strong>Article References:</strong> Rangesh, K., Rajan, R. S. P., Dineshbabu, M., Dhayalan, R. E., Johnson Arun Kumar, C., Tharmadurai, S., Anand, M., &amp; Panda, U. S. (2026). Aragonite saturation state indicates emerging ocean acidification hotspots in the Gulf of Mannar and Palk Bay along the Southeast Coast of India. <em>Discover Oceans, 3</em>(1), Article 52. <a href="https://doi.org/10.1007/s44289-026-00165-x" rel="noopener noreferrer">https://doi.org/10.1007/s44289-026-00165-x</a></p>
<p><strong>Image Credits:</strong> AI Generated</p>
<p><strong>DOI:</strong> <a href="https://doi.org/10.1007/s44289-026-00165-x" rel="noopener noreferrer">10.1007/s44289-026-00165-x</a></p>
<p><strong>Keywords:</strong> ocean acidification, aragonite saturation, Gulf of Mannar, Palk Bay, coral reefs, seagrass meadows, carbonate chemistry, Indian Ocean, Aragonite, saturation, state, indicates</p>
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