\\n\\n
Dr. Pletser’s experience includes 30 years of working with the European Space Agency as a Senior Physicist/Engineer and coordinating their parabolic flight campaigns, and he is the Guinness World Record holder for the most number of aircraft flown (12) in parabolas, personally logging more than 7,300 parabolas.
\\n\\nSeeing the 5,000th book published makes us at the same time proud, happy, humble, and grateful. This is a great opportunity to stop and celebrate what we have done so far, but is also an opportunity to engage even more, grow, and succeed. It wouldn't be possible to get here without the synergy of team members’ hard work and authors and editors who devote time and their expertise into Open Access book publishing with us.
\\n\\nOver these years, we have gone from pioneering the scientific Open Access book publishing field to being the world’s largest Open Access book publisher. Nonetheless, our vision has remained the same: to meet the challenges of making relevant knowledge available to the worldwide community under the Open Access model.
\\n\\nWe are excited about the present, and we look forward to sharing many more successes in the future.
\\n\\nThank you all for being part of the journey. 5,000 times thank you!
\\n\\nNow with 5,000 titles available Open Access, which one will you read next?
\\n\\nRead, share and download for free: https://www.intechopen.com/books
\\n\\n\\n\\n
\\n"}]',published:!0,mainMedia:null},components:[{type:"htmlEditorComponent",content:'
Preparation of Space Experiments edited by international leading expert Dr. Vladimir Pletser, Director of Space Training Operations at Blue Abyss is the 5,000th Open Access book published by IntechOpen and our milestone publication!
\n\n"This book presents some of the current trends in space microgravity research. The eleven chapters introduce various facets of space research in physical sciences, human physiology and technology developed using the microgravity environment not only to improve our fundamental understanding in these domains but also to adapt this new knowledge for application on earth." says the editor. Listen what else Dr. Pletser has to say...
\n\n\n\nDr. Pletser’s experience includes 30 years of working with the European Space Agency as a Senior Physicist/Engineer and coordinating their parabolic flight campaigns, and he is the Guinness World Record holder for the most number of aircraft flown (12) in parabolas, personally logging more than 7,300 parabolas.
\n\nSeeing the 5,000th book published makes us at the same time proud, happy, humble, and grateful. This is a great opportunity to stop and celebrate what we have done so far, but is also an opportunity to engage even more, grow, and succeed. It wouldn't be possible to get here without the synergy of team members’ hard work and authors and editors who devote time and their expertise into Open Access book publishing with us.
\n\nOver these years, we have gone from pioneering the scientific Open Access book publishing field to being the world’s largest Open Access book publisher. Nonetheless, our vision has remained the same: to meet the challenges of making relevant knowledge available to the worldwide community under the Open Access model.
\n\nWe are excited about the present, and we look forward to sharing many more successes in the future.
\n\nThank you all for being part of the journey. 5,000 times thank you!
\n\nNow with 5,000 titles available Open Access, which one will you read next?
\n\nRead, share and download for free: https://www.intechopen.com/books
\n\n\n\n
\n'}],latestNews:[{slug:"stanford-university-identifies-top-2-scientists-over-1-000-are-intechopen-authors-and-editors-20210122",title:"Stanford University Identifies Top 2% Scientists, Over 1,000 are IntechOpen Authors and Editors"},{slug:"intechopen-authors-included-in-the-highly-cited-researchers-list-for-2020-20210121",title:"IntechOpen Authors Included in the Highly Cited Researchers List for 2020"},{slug:"intechopen-maintains-position-as-the-world-s-largest-oa-book-publisher-20201218",title:"IntechOpen Maintains Position as the World’s Largest OA Book Publisher"},{slug:"all-intechopen-books-available-on-perlego-20201215",title:"All IntechOpen Books Available on Perlego"},{slug:"oiv-awards-recognizes-intechopen-s-editors-20201127",title:"OIV Awards Recognizes IntechOpen's Editors"},{slug:"intechopen-joins-crossref-s-initiative-for-open-abstracts-i4oa-to-boost-the-discovery-of-research-20201005",title:"IntechOpen joins Crossref's Initiative for Open Abstracts (I4OA) to Boost the Discovery of Research"},{slug:"intechopen-hits-milestone-5-000-open-access-books-published-20200908",title:"IntechOpen hits milestone: 5,000 Open Access books published!"},{slug:"intechopen-books-hosted-on-the-mathworks-book-program-20200819",title:"IntechOpen Books Hosted on the MathWorks Book Program"}]},book:{item:{type:"book",id:"805",leadTitle:null,fullTitle:"Sex Steroids",title:"Sex Steroids",subtitle:null,reviewType:"peer-reviewed",abstract:'This book, entitled "Sex Steroids", features a valuable collection of reviews and research articles written by experts in signal transduction, cellular biology, diseases and disorders. 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by"}}],ofsBooks:[]},correction:{item:{id:"66063",slug:"corrigendum-to-introductory-chapter-historical-perspective-and-brief-overview-of-insulin",title:"Corrigendum to: Introductory Chapter: Historical Perspective and Brief Overview of Insulin",doi:null,correctionPDFUrl:"https://cdn.intechopen.com/pdfs/66063.pdf",downloadPdfUrl:"/chapter/pdf-download/66063",previewPdfUrl:"/chapter/pdf-preview/66063",totalDownloads:null,totalCrossrefCites:null,bibtexUrl:"/chapter/bibtex/66063",risUrl:"/chapter/ris/66063",chapter:{id:"63640",slug:"introductory-chapter-historical-perspective-and-brief-overview-of-insulin",signatures:"Gaffar Sarwar Zaman",dateSubmitted:"June 29th 2018",dateReviewed:"August 28th 2018",datePrePublished:"November 5th 2018",datePublished:"February 6th 2019",book:{id:"6675",title:"Ultimate Guide to Insulin",subtitle:null,fullTitle:"Ultimate Guide to Insulin",slug:"ultimate-guide-to-insulin",publishedDate:"February 6th 2019",bookSignature:"Gaffar 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Insulin",subtitle:null,fullTitle:"Ultimate Guide to Insulin",slug:"ultimate-guide-to-insulin",publishedDate:"February 6th 2019",bookSignature:"Gaffar Zaman",coverURL:"https://cdn.intechopen.com/books/images_new/6675.jpg",licenceType:"CC BY 3.0",editedByType:"Edited by",editors:[{id:"203015",title:"Dr.",name:"Gaffar Sarwar",middleName:"Sarwar",surname:"Zaman",slug:"gaffar-sarwar-zaman",fullName:"Gaffar Sarwar Zaman"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}},authors:[{id:"203015",title:"Dr.",name:"Gaffar Sarwar",middleName:"Sarwar",surname:"Zaman",fullName:"Gaffar Sarwar Zaman",slug:"gaffar-sarwar-zaman",email:"gffrzaman@gmail.com",position:null,institution:{name:"King Khalid University",institutionURL:null,country:{name:"Saudi Arabia"}}}]},book:{id:"6675",title:"Ultimate Guide to Insulin",subtitle:null,fullTitle:"Ultimate Guide to Insulin",slug:"ultimate-guide-to-insulin",publishedDate:"February 6th 2019",bookSignature:"Gaffar Zaman",coverURL:"https://cdn.intechopen.com/books/images_new/6675.jpg",licenceType:"CC BY 3.0",editedByType:"Edited by",editors:[{id:"203015",title:"Dr.",name:"Gaffar Sarwar",middleName:"Sarwar",surname:"Zaman",slug:"gaffar-sarwar-zaman",fullName:"Gaffar Sarwar Zaman"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}}},ofsBook:{item:{type:"book",id:"9484",leadTitle:null,title:"Glucagon",subtitle:null,reviewType:"peer-reviewed",abstract:'
\r\n\tGlucagon is one of the pancreatic hormones. It plays different and significant roles in the human body. Their functions include protections against damages of glucose homeostasis, in particular against hypoglycemia. It influences and regulates not only metabolism and glucose homeostasis. Glucagon is also involved in the metabolism of lipids and amino acids, as well as it regulates also energy homeostasis. On the other hand, disturbances in functions, synthesis and/or secretion of glucagon may cause several pathologies such as obesity and hepatic steatosis. Impairment functions of this hormone may have influence, direct or indirect, on the development of diabetes mellitus, "a progressive worldwide epidemic". Presented examples of glucagon\'s role in human health and diseases suggest a very important role of this hormone. Therefore, knowledge of this hormone, its functions, regulations, secretions, etc seems to be very significant. This book will be written by a scientist with expertise in the study of glucagon.
\r\n\r\n\tThis book describes a series of up-to-date topics about physiological and pathological processed that occurred in the functions of glucagon. By presenting a clear and exhaustive review of the correlation between synthesis and secretion of glucagon and health/disease, it is expected to draw more attention from biomedical scientists, physicians, pharmacologists, physiologists, and students to dedicate their research in uncovering the role of hormonal regulation, especially glucagon, in human health and diseases.
',isbn:null,printIsbn:"979-953-307-X-X",pdfIsbn:null,doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!1,hash:"e2fce59d7d576c75b80cbdd2e271fe54",bookSignature:"Dr. Leszek Szablewski",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/9484.jpg",keywords:"Pancreatic Hormones, Glucagon and Glucose Homeostasis, Regulation of Glucagon Synthesis, Regulation of Glucagon Secretion, Gluconeogenesis, Glycogenolysis, Characteristics of Glucagon, Glucagon Signaling Pathway, Hyperglucagonemia, Glucagonoma, Glucagon Resistance, Hepatic Steatosis",numberOfDownloads:null,numberOfWosCitations:0,numberOfCrossrefCitations:0,numberOfDimensionsCitations:0,numberOfTotalCitations:0,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"October 30th 2019",dateEndSecondStepPublish:"March 27th 2020",dateEndThirdStepPublish:"May 26th 2020",dateEndFourthStepPublish:"August 14th 2020",dateEndFifthStepPublish:"October 13th 2020",remainingDaysToSecondStep:"a year",secondStepPassed:!0,currentStepOfPublishingProcess:5,editedByType:null,kuFlag:!1,biosketch:null,coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"49739",title:"Dr.",name:"Leszek",middleName:null,surname:"Szablewski",slug:"leszek-szablewski",fullName:"Leszek Szablewski",profilePictureURL:"https://mts.intechopen.com/storage/users/49739/images/system/49739.jpg",biography:"Leszek Szablewski is a professor of medical sciences. He received his M.S. in the Faculty of Biology from the University of Warsaw and his PhD degree from the Institute of Experimental Biology Polish Academy of Sciences. He habilitated in the Medical University of Warsaw, and he obtained his degree of Professor from the President of Poland. Professor Szablewski is the Head of Chair and Department of General Biology and Parasitology, Medical University of Warsaw. Professor Szablewski has published over 80 peer-reviewed papers in journals such as Journal of Alzheimer’s Disease, Biochim. Biophys. Acta Reviews of Cancer, Biol. Chem., J. Biomed. Sci., and Diabetes/Metabol. Res. Rev, Endocrine. He is the author of two books and four book chapters. He has edited four books, written 15 scripts for students, is the ad hoc reviewer of over 30 peer-reviewed journals, and editorial member of peer-reviewed journals. Prof. Szablewski’s research focuses on cell physiology, genetics, and pathophysiology. He works on the damage caused by lack of glucose homeostasis and changes in the expression and/or function of glucose transporters due to various diseases. 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From chapter submission and review, to approval and revision, copyediting and design, until final publication, I work closely with authors and editors to ensure a simple and easy publishing process. I maintain constant and effective communication with authors, editors and reviewers, which allows for a level of personal support that enables contributors to fully commit and concentrate on the chapters they are writing, editing, or reviewing. I assist authors in the preparation of their full chapter submissions and track important deadlines and ensure they are met. I help to coordinate internal processes such as linguistic review, and monitor the technical aspects of the process. As an ASM I am also involved in the acquisition of editors. 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Mauricio Barría",coverURL:"https://cdn.intechopen.com/books/images_new/6550.jpg",editedByType:"Edited by",editors:[{id:"88861",title:"Dr.",name:"R. Mauricio",surname:"Barría",slug:"r.-mauricio-barria",fullName:"R. Mauricio Barría"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"1591",title:"Infrared Spectroscopy",subtitle:"Materials Science, Engineering and Technology",isOpenForSubmission:!1,hash:"99b4b7b71a8caeb693ed762b40b017f4",slug:"infrared-spectroscopy-materials-science-engineering-and-technology",bookSignature:"Theophile Theophanides",coverURL:"https://cdn.intechopen.com/books/images_new/1591.jpg",editedByType:"Edited by",editors:[{id:"37194",title:"Dr.",name:"Theophanides",surname:"Theophile",slug:"theophanides-theophile",fullName:"Theophanides Theophile"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"3092",title:"Anopheles mosquitoes",subtitle:"New insights into malaria vectors",isOpenForSubmission:!1,hash:"c9e622485316d5e296288bf24d2b0d64",slug:"anopheles-mosquitoes-new-insights-into-malaria-vectors",bookSignature:"Sylvie Manguin",coverURL:"https://cdn.intechopen.com/books/images_new/3092.jpg",editedByType:"Edited by",editors:[{id:"50017",title:"Prof.",name:"Sylvie",surname:"Manguin",slug:"sylvie-manguin",fullName:"Sylvie Manguin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"3161",title:"Frontiers in Guided Wave Optics and Optoelectronics",subtitle:null,isOpenForSubmission:!1,hash:"deb44e9c99f82bbce1083abea743146c",slug:"frontiers-in-guided-wave-optics-and-optoelectronics",bookSignature:"Bishnu Pal",coverURL:"https://cdn.intechopen.com/books/images_new/3161.jpg",editedByType:"Edited by",editors:[{id:"4782",title:"Prof.",name:"Bishnu",surname:"Pal",slug:"bishnu-pal",fullName:"Bishnu Pal"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"72",title:"Ionic Liquids",subtitle:"Theory, Properties, New Approaches",isOpenForSubmission:!1,hash:"d94ffa3cfa10505e3b1d676d46fcd3f5",slug:"ionic-liquids-theory-properties-new-approaches",bookSignature:"Alexander Kokorin",coverURL:"https://cdn.intechopen.com/books/images_new/72.jpg",editedByType:"Edited by",editors:[{id:"19816",title:"Prof.",name:"Alexander",surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"1373",title:"Ionic Liquids",subtitle:"Applications and Perspectives",isOpenForSubmission:!1,hash:"5e9ae5ae9167cde4b344e499a792c41c",slug:"ionic-liquids-applications-and-perspectives",bookSignature:"Alexander Kokorin",coverURL:"https://cdn.intechopen.com/books/images_new/1373.jpg",editedByType:"Edited by",editors:[{id:"19816",title:"Prof.",name:"Alexander",surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"55299",title:"An Insightful Model to Study Innate Immunity and Stress Response in Deep‐Sea Vent Animals: Profiling the Mussel Bathymodiolus azoricus",doi:"10.5772/68034",slug:"an-insightful-model-to-study-innate-immunity-and-stress-response-in-deep-sea-vent-animals-profiling-",body:'\nDeep‐sea hydrothermal vents were discovered 40 years ago in the Galapagos Rift [1] revealing for the first time, to the amazement of the scientific community, unusual life forms that have developed unique biochemical adaptations to high temperatures and toxic chemical nature of vent surrounding, otherwise harmful to life as we know it on the surface of the planet [2–5]. The animals dwelling around the vent sites exhibit high productivity and thus must cope with the seemingly deleterious physical and chemical conditions, while developing surprisingly successful strategies to withstand adverse environmental conditions, including environmental microbes and mechanical stress whether due to animal predation or from deep‐sea volcanic eruptions [6–8].
\nAt such depths and in the absence of light, life is thriving in chemosynthesis‐based ecosystems where most abundant marine invertebrates have developed mutualistic relationships with chemosynthetic bacteria. These symbiotic interactions are believed to play a crucial role in the survival of hydrothermal vent animals, driving their transcriptional activities, and their successful adaptation strategies to subsist under extreme environmental conditions. They essentially rely on the establishment of endosymbiosis relationships between vent animals and sulfur‐oxidizing (SOX) or methane‐oxidizing (MOX) bacteria [9–13].
\nDeep‐sea vent mussels of the Bathymodiolus genus are dominant members at hydrothermal vents and cold seep habitats. These mussels have the peculiarity of sheltering both endosymbiotic sulfide‐oxidizing and methane‐oxidizing bacteria in their gills [9–13], supporting thus their endurance within this type of environment. Bathymodiolus azoricus is also the dominant species in deep‐sea hydrothermal vents in the Azores region and is well adapted to extreme conditions particularly to toxic concentrations of heavy metals, acidic pH, and absence of light [14–17].
\nIn an attempt to understand physiological reactions of animals normally set to endure extreme conditions, in deep‐sea environments, our laboratory has undertaken, for the last 6 years, a series of investigations aimed at characterizing molecular indicators of adaptation processes of which components of the immune and stress‐related systems have received most of our attention [18, 19]. Central to our studies is the long‐term maintenance of vent mussels to atmospheric pressure proven to be a useful model to study unique molecular relationships under which the regulation of gene transcription may be affected by aquaria conditions and by the gradual disappearance of endosymbiont bacteria from gill epithelia [20]. Nonetheless, vent mussels subsist for months at atmospheric pressure in aquaria supplemented with plain sea water in or with artificial diet. This has allowed us to focus on developing experiments to investigate new physiological responses of animals sustaining experimental challenges involving immunological and stress‐related reactions and to provide new approaches to assess the effect of natural microorganisms and metal toxicity at vent environments [21, 22]. As a research model, the choice of the vent mussel B. azoricus is of great significance given its unique symbiosis with SOX and MOX bacteria. It has provided us with the means to understanding the molecular mechanisms underlying immune reactions in animals normally set to endure extreme deep‐sea environments and the role of their symbiotic bacteria in controlling immune gene transcriptional activity.
\nIn line with this, we have investigated main constituents of the vent mussel immune system and demonstrated how immune and stress genes could be modulated upon different experimental challenges in the absence of the characteristic high hydrostatic pressure found at deep‐sea vent sites without methane and/or sulfide supplementation [21–23]. The proximity to the nearby hydrothermal vent fields, in the Azores region, has given us a geographical advantage for earning first insight into immediate physiological responses comprising both cellular and humoral responses of live mussels, freshly collected from the hydrothermal vents, which upon arrival, are acclimatized to our aquarium system, LabHorta [23–25]. The maintenance of live mussels in our laboratory is thus a key factor in gaining knowledge into the physiology of vent animals including the study of evolutionary conserved immune, inflammatory, and stress‐related factors commonly found in other marine bivalves [19, 26].
\nThe interaction between microorganisms and host defense mechanisms is a decisive factor for the survival of marine bivalves. They rely on cell‐mediated and humoral reactions to overcome the pathogens that naturally occur in the marine environment [27]. Growing interest in deep‐sea vent biology has turned the vent mussel B. azoricus into a model organism centered on research activities based on the premise that vent mussels clearly have need for an immune system to overcome microbial challenges in their natural surroundings. For this reason, our research strategies have been focused on the molecular characterization of molecules participating in immune reactions, using in vivo and ex vitro models, to elucidate cellular and humoral defense mechanisms in vent mussels and its survival strategies under extreme environments. As for other bivalves, the innate immune system of B. azoricus is based on cellular constituents and soluble hemolymph (blood) factors, which play a prominent role in protecting the animals against invading microorganisms. The circulating hemocytes or blood cells are mostly found in the hemolymph and extrapallial fluid. They are responsible for cell‐mediated defense reactions such as phagocytosis and the activation of a variety of cytotoxic reactions including the release of lysozomal enzymes and antimicrobial peptides [28–30]. Moreover, the generation of highly reactive oxygen intermediates (ROIs) and nitric oxide also plays an important defense role against pathogens [30–33]. Besides their decisive role in protecting the host from microbial assaults, bivalve hemocytes have also been implicated in other important physiological functions, including nutrient transport, digestion, wound healing and shell regeneration and/or mineralization, and excretion [34]. In addition, the hemolymph serum contains humoral defense factors such as lectins and cytokine‐like molecules that are directly and indirectly involved in the killing of pathogens and in mediating cell‐cell interactions, respectively. Lectins are important mediators of cellular reactions and exhibit opsonin properties, which facilitate the phagocytosis [35–39]. The hemolymph also contains antibacterial factors and lysozomal components that ensure, along with hemocyte phagocytic and cytotoxic processes, the clearance of pathogenic bacteria [38, 39]. Using a combination of light microscopy and staining procedures, three major hemocyte types are discernible in the extrapallial fluid and hemolymph of B. azoricus. The most abundant type was identified as granulocyte readily recognizable by their cytoplasmic granules [19]. They appear fairly homogeneous in size and showing a characteristic crescent, or half‐moon shape morphology upon adherence to glass slides and before migratory movements. Granulocytes spread well onto the glass surface averaging 30–40 μm in length. In contrast, hyalinocytes presented smoother cytoplasm, i.e., a nongranular appearance due to a lower amount of cytoplasmic granules noticeable under phase contrast and differential interference contrast visualizations [19]. A third less common hemocyte type was also observed. They correspond to hemoblast‐like cells and presented a spherical shape appearance with higher nucleus to cytoplasm ratio when compared to granulocytes and hyalinocytes [19]. In vitro phagocytic assays carried out with B. azoricus hemocytes revealed that 70% of the hemocytes containing more than two zymosan particles were granulocytes and to a lesser extent the percentage of phagocytic cells corresponding to hyalinocytes was 23%. In contrast, the percentage of hemoblasts containing ingested zymosan particles was 5–7%, the lowest revealed in our studies [19].
\nAlong with hemocytes studies, we began to tackle signaling pathways putatively involved in the mediation of cellular responses in the presence of Vibrio spp. It was demonstrated that compounds of microbial origin could trigger detectable phosphorylation events in B. azoricus hemocyte extracts and likely involving the activation of different classes of mitogen‐activated protein kinases (MAPKs). When challenged with a marine bacterium, Vibrio parahaemolyticus or a nonmarine bacterium, Bacillus subtilis, to stimulate hemocytes, cellular proteins were differently phosphorylated as demonstrated in Western blotting experiments using the MAPK/ERK, p38, and JNK rabbit polyclonal antibodies. Moreover, the differences seen in phosphorylation patterns could be attributed to inherent properties of the bacterial strain used, differences in the mechanisms of binding to hemocytes, or differential activation of cell membrane receptors and signaling pathways, resulting in different patterns of protein phosphorylation. Western blotting analyses suggest that B. azoricus hemocytes display receptors with binding affinities toward microbial molecules or to live bacteria [19].
\nAs mytilid species, the deep‐sea vent mussel B. azoricus exhibits large lamellated gills (ctenidia) arranged as numerous filament structures stacked together through ciliary junctions. Each filament is organized in two coalescent epithelial cell sheets overlaying a central lumen where hemocytes can be found. The thinness of gill filaments allows for the visualization of live hemocytes through the epithelium, and to monitor hemocyte motility directly under light microscopy [19]. Bivalve mollusk gills assume thus a strategic importance at the interface between the external milieu and the internal body cavities of the animal where contact with microorganisms is inevitable during feeding processes inasmuch as host defense responses may incur from interactions with infective pathogens during normal filtration [36–39]. For this reason, a number of typical cellular and humoral immune reactions are likely to take place in gill tissues and observable as hemocyte proliferation and phagocytosis, the activation of immune signaling pathways, and the activation of genes involved in immune, antioxidant and antibacterial responses against invading bacterial pathogens or the presence of metal toxicants [36–39].
\nTo further test the gill’s ability to mount immune reactions, a series of ex vivo experiments have been performed using gill tissues freshly dissected from vent mussels and subjected to short‐term incubations in tissue culture well‐plates and under different experimental settings. Different stimuli were carried out to demonstrate the expression of genes in gill tissues exposed to a mixture of endosymbionts previously obtained from gill extracts, and V. parahaemolyticus, in comparison with sterile sea water incubations (Figure 1). Differential gene expression results indicated that exposure to methanotrophic and thiotrophic endosymbiont preparations led to general upregulation of genes involved in immune recognition reactions, without the addition of hemolymph, while gill incubations with endosymbiont extract and to which hemolymph serum (hemocyte free) was added led to an opposite effect, resulting in a lower expression of immune recognition genes. These results contrasted with incubations performed with V. parahaemolyticus or with control sterile sea water. In this case, higher levels of gene expression were achieved when hemolymph was added to gill tissues incubated with V. parahaemolyticus or control sterile sea water (Figure 1). Ex vivo experiments as described bring evidence supporting a yet uncharacterized effect of hemolymph and its humoral constituents over endosymbionts, likely controlling immune gene expression of its host B. azoricus. This prompted the question of whether the host gill tissue would be able or not to recognize endosymbiont as self‐particles and to which extent the host immune system does not disturb the acquisition of endosymbionts by horizontal transfer during the host larval stages. Moreover, the permanence and survival of endosymbionts within gill tissue would require a fitted control over the host immune system, acting on the transcriptional regulation of immune genes and at the level of pattern recognition receptors (PRRs) expressed by cells of the host innate immune system to detect microbial‐associated molecular patterns (MAMPs) present on the surface of microorganisms [40, 41]. This endosymbiont effect over the host immune system would likely require the presence of hemolymph and its humoral constituents, as demonstrated by the gill ex vivo experiments (Figures 1 and 2). Since the gill tissue does grow over the mussel life’s time we also probed different gill sections to determine levels of immune gene expression along the anterior‐posterior axis, notably the “budding zone” on the posterior end, considered as the youngest section of the gill and through which endosymbionts are believed to make their entry. It was found that vent mussel gene expressions were markedly lower than other gill tissue sections, suggesting a thigh mechanism of transcriptional regulation of host genes in the presence of endosymbiont bacteria in the gill budding zone.
\nEx vivo experiments performed with dissected gills. Gill fragments were incubated with V. parahaemolyticus and with an enriched preparation of endosymbionts freshly obtained from gill homogenates and gradient centrifugation. The effect of hemolymph humoral factors was tested by incubating gill fragments with and without hemolymph in the presence of V. parahaemolyticus and endosymbiont mixture. Results were compared to control incubations with plain sterile sea water. Gene expression was performed by qPCR targeting the immune recognition genes Aggrecan, C‐type lectin, C‐type lectin immune receptor, immune lectin receptor, lipopolysaccharide‐binding protein‐bactericidal/permeability‐increasing protein, peptidoglycan recognition protein, and serine protease inhibitor‐2 [50]. Incubations performed with endosymbiont preparations distinctively induced immune recognition genes in the absence of hemocyte‐free hemolymph.
Ex vivo experiments performed with dissected gills. As in Figure 1, gill fragments were incubated with distinct bacterial stimulants: an enriched methanotrophic bacterial preparation; an enriched thiotrophic bacterial preparation, a mixture of methanotrophic and thiotrophic bacterial preparations, Vibrio parahaemolyticus and Photobacteria bacterium. Results were compared to control incubations with plain sterile sea water. Gene expression was performed by qPCR targeting same immune recognition genes as in Figure 1. Separate methanotrophs and thiotrophs preparations induced higher levels of immune gene expressions when compared to a mixture of the two endosymbiont bacteria preparations. Incubations using Vibrio parahaemolyticus resulted in drastic downregulation of immune recognition genes.
As filter feeders living most of their lives attached to a substrate, bivalves are exposed to constant biologically available pollutants over an extended period of time [42–44]. They have been studied as biological models to assess the impact of pollution in the environment and used as a biomonitoring “tool” due to their capacity of bioaccumulating high concentrations of trace metals, mostly in soft tissues such as gills and digestive gland [45–47]. The large surface of the gills and their involvement in gas exchange and feeding processes bring bivalves to constant and intimate contact with their environment where pathogens may also find their route of entry and encounter the bivalve first‐line immune defense reactions.
\nWhile marine bivalves living in sandy, rocky intertidal, and shallow subtidal environments may rely on well‐established humoral and cellular immune reactions to counteract pathogenic microorganisms, a new level of molecular intricacy may be seen between endosymbiont‐bearing bivalves living in anaerobic and sulfide‐rich environments and the pathogenic microorganisms they encounter. These natural molecular interactions would account for the role of endosymbionts in modulating the host immunity by controlling the transcriptional activity of immune genes. Bivalve associations with chemoautotrophic endosymbionts are now well known and widely distributed across a range of different chemosynthetic environments, including deep‐sea hydrothermal vents (Bathymodiolus spp., Calyptogena sp.); gas seeps, mud volcanoes, and petroleum seeps (Bathymodiolus spp., Calyptogena sp.); whale and wood falls (Idas spp., Adipicola spp., Vesicomya sp., Axinodon sp.); and shallow water anoxic sediments mediated by sulfate reduction (Solemya spp., Codaki spp., Anodontia spp., Lucina spp.) [7].
\nOur recent results from ex vivo gill tissue experiment proven to be a valuable system for the study of tissue‐specific immune responses where the thin epithelial cell layers of gill filaments would make it possible to signal pathogen‐sensing directly through gill epithelia and affecting adjacent methanotrophic or thiotrophic endosymbionts which in turn would functionally prime host immune cells, the hemocytes, into altering their transcriptional activity (Figure 3). The endosymbiont immunomodulatory effect on the host immune system, as discussed in detail further below, is still under current investigations by our research team as the complexity of host‐endosymbiont interactions in the deep‐sea vent mussel B. azoricus remains to be fully understood. This model is consistent with the hypothesis that innate immune receptors are required to promote long‐term colonization by microbiota. This emerging perspective challenges current paradigms in immunology and suggests that PRRs may have evolved, in part, to mediate the bidirectional cross‐talk between microbial symbionts and their hosts [48, 49].
\nHypothetical model representing the host‐endosymbiont‐mediated immune responses against pathogens. In a normal immunological state, hemocytes PRRs are being sensitized by host‐endosymbiont interactions allowing the vent mussel immune system to remain active and tolerant to the presence of MOX and SOX bacteria. Upon interacting with extracellular pathogens, host‐symbiont interactions are altered and incur in higher endosymbiont genes transcriptional activity [74] and subsequently affecting host hemocytes by triggering its immune repertoire via PRRs activation.
In 2010 a high‐throughput sequencing and analysis of the gill tissue transcriptome from the deep‐sea hydrothermal vent mussel B. azoricus was reported by our group [50]. It represented the first tissue transcriptional analysis of a deep‐sea hydrothermal vent animal, using next generation sequencing technology, enabling the creation of a searchable catalog of genes that provided a direct method of identifying and retrieving vast numbers of novel coding sequences which could then be applied in gene expression profiling experiments, using quantitative polymerase chain reaction (qPCR), from a nonconventional model organism [50]. It provided the most comprehensive sequence resource for identifying novel genes currently available for a deep‐sea vent organism, in particular, genes putatively involved in immune and inflammatory reactions in vent mussels. This first transcriptional analysis of gill tissues from the deep‐sea hydrothermal vent B. azoricus was organized as a searchable catalog of genes providing a direct method of identifying and retrieving vast numbers of novel coding sequences, which can be applied in gene expression profiling experiments. The assembled and annotated sequences were organized in a dedicated database, accessible through the website
With an unprecedented high number of gene sequences available from our transcriptomic data, we were able to tackle signaling pathways and compare gene expression profiles in a series of experiments aiming at better understanding innate immunity in animals physiologically programmed to endure deep‐sea vent conditions. Responses to bacterial infections with different strains of Vibrio wound experiments, long‐term acclimatization in aquarium conditions and pressurization experiments with the hyperbaric chamber IPOCAMP [51] became the main focus of our research, setting thus the grounds for more in‐depth analyzes revealing distinct gene expression profiles behind unique molecular relationships under which the regulation of gene transcription may be affected by biotic factors including microorganisms, the presence of endosymbiont bacteria and shell damage incurring in opportunistic infections or by abiotic factor as the hydrostatic pressure. The majority of the genes comprising four functional categories as described by Bettencourt et al. [50] and relating to immune recognition, signaling transduction, transcription, and effector molecules mechanisms were analyzed by qPCR.
\nThe long‐term aquarium maintenance of vent mussels to atmospheric pressure has long been central to our studies and proven to be a useful model to study unique molecular relationships under which the regulation of gene transcription may be affected by the gradual disappearance of endosymbiont bacteria from gill epithelia [20, 52]. Nonetheless, vent mussels from Menez Gwen hydrothermal vent site subsist for months at atmospheric pressure in aquarium conditions, in plain sea water or supplemented with methane and sulfide. This has allowed us to focus on developing experiments to investigate new physiological responses of vent mussels sustaining experimental challenges involving bacterial pathogens of the Vibrio genus, even in the absence of the characteristic high hydrostatic pressure found at deep‐sea vent sites and without methane and sulfide supplementation [21, 22].
\nEarlier results from experimental exposures to Vibrio splendidus, Vibrio alginolyticus, Vibro anguillarum, and Flavobacterium sp. pointed at the immune discriminatory capacity of B. azoricus to distinguish different Vibrio strains, and at significant differences of immune gene expression levels between 12 and 24 h exposure times. These studies concluded that the immune gene transcriptional activity was modulated at two levels, i.e., over the course of time and according to the bacterial strain tested, suggesting thus, a selective response toward Vibrio spp. when vent mussels were experimentally challenged during 24 h [53, 54]. Additional experiments were carried out with Vibrio diabolicus aiming at the analysis of gene expression differences between distinct vent mussel populations from the hydrothermal vent sites Menez Gwen (MG, 800 m depth) and Lucky Strike (LS, 1700 m depth) both located on the Mid‐Atlantic region, near the Azores islands. These comparative studies revealed unique immune transcriptional specificities at the gill, digestive gland, and mantle tissues level providing further evidence supporting different usage of transcription factors at the promoter region of immune genes possibly linked to the hydrothermal vent environment Furthermore, Menez Gwen (MG) and Lucky Strike (LS) B. azoricus showed significant gene expression differences during V. diabolicus challenges over time demonstrating that immune genes are differentially expressed within the same mussel populations regardless of their hydrothermal vent origin suggesting thus site‐related tissue‐specific gene expression patterns [55]. Moreover, these results also suggested different tissue tolerance to decompression and adaptation to atmospheric pressure not seen so far. Mantle tissues from LS mussels seemed unaffected by deep‐sea retrieval showing significantly higher levels of immune gene expressions as compared to MG mantle tissues. Thus, the decompression effect on the animal’s internal organs may be evaluated by ways of its ability to respond, at the immune transcriptional level, to V. diabolicus challenges. For that reason, mantle tissues from LS animals appear to be decompression‐resistant and immune competent toward bacterial challenges. On the other hand, the digestive gland revealed the most increased gene expression levels in MG animals showing how the tissue microenvironment is relevant to in situ immune responses. Gill immune transcriptional activity in both MG and LS mussels was not as significantly different as for the other tissues tested which may be attributed to the presence of endosymbiont bacteria in gill epithelia acting as a driving factor likely to affect host‐gene expression and the overall physiological statuses of MG and LS vent mussels while interacting with V. diabolicus. Even though gill tissues have been the main focus of most of our previous investigations in the deep‐sea vent mussel B. azoricus, the digestive gland and mantle tissues hold the potential for highlighting specific immune responses in tissues other than gills and how they can modulate the outcome of the animal’s overall immune responses [55].
\nIn addition to ex vivo experiments and Vibrio exposures to live vent mussels, we were able to carry out long‐term acclimatization experiments with vent mussels kept in aquaria and at atmospheric pressure. These experiments were devised to assess the effect of such prolonged aquarium conditions on immune and stress‐related reactions as mussels were gradually releasing their endosymbiont bacteria from gill bacteriocytes. These studies provided a basis for understanding the interactions between host‐immune and endosymbiont gene expressions during postcapture long‐term acclimatization in plain sea water and represented an ideal model for investigating B. azoricus immune genes transcriptional activity and symbiont bacteria prevalence, in view of changes in the availability of chemical‐based energy sources during acclimatization at atmospheric pressure. It also pointed out the relevance of gene expression studies while addressing the swift changes affecting metabolic adaptations and food intake fluctuations, whether induced by or as a result of the gradual loss of endosymbionts and subsequent presence of symbiont bacteria in the aquarium environment, altering thus the physiological homeostasis of B. azoricus [56]. These studies demonstrated that the transcriptional activity profiles for immune and bacterial endosymbiont genes followed a time‐dependent mRNA transcriptional pattern evidenced at 24 h, 1 week, and 3 weeks acclimatization. Furthermore, after 1 week acclimatization, vent mussels were under the influence of what appears to be a concomitant host‐immune and endosymbiont gene expression, possibly indicating a physiological transition point which induces higher levels of transcriptional activity [56]. Under such circumstances, survival of vent mussels will require immune gene repertoire switching involving the differential expression (DE) of recognition, signaling, transcription, and effector genes tied to environmental parameters and to the symbiotic relationships in B. azoricus. Metabolic adaptations and food intake changes, whether induced as a result of the gradual loss of endosymbionts and subsequent release in the aquarium environment, are likely to affect gene transcription activities and prevalence of symbionts in gill tissues [56–58].
\nThe geographic proximity to the nearby hydrothermal vent fields, in the Azores region, gave our laboratory a positional advantage for earning first insight into immediate physiological responses comprising both cellular and humoral responses of freshly collected mussels from different hydrothermal vents, which upon arrival, are acclimatized to our aquarium system, LabHorta [23].The maintenance of live mussels from the shallower vent field, Menez Gwen, became thus a key factor in gaining knowledge into the physiology of vent animals including the study of evolutionary conserved immune, inflammatory and stress‐related factors commonly found in other marine bivalves [18–22].
\nTaking advantage of the LabHorta facility, comparisons studies were made possible, with live vent mussels subjected to V. parahaemolyticus infection, wound injury, hyperbaric pressurization, and 3 months acclimatization (Figure 4). These experiments allowed for the characterization of the differential activation of signaling pathways and the relative quantification of immune genes expressed during each type of stimulation. Differential gene expression results indicated that the four experimental conditions tested were distinctively inducing the immune genes of vent mussels to different levels of transcriptional activity of which the immune and signal transduction genes showed the highest expressions (Figure 5).
\nSchematic representation of immune signaling activation. After initial events characterized by immune recognition and stress‐related reactions, signal transduction pathways are induced into transmitting a series of protein phosphorylation events, through the intracellular milieu, which ultimately result in the translocation of transcription factors into the nucleus that initiates the transcription of genes encoding immune effector molecules, here represented as lysozyme, metallothionein, and ferritin.
Comparative gene expression profiles from vent mussels subjected to 3 months acclimatization in aquaria at atmospheric pressure; Vibrio parahaemolyticus exposures; wound injury, and repressurization in the IPOCAMP chamber. Results are presented as relative expression folds calculated by qPCR and targeting immune genes from recognition, signaling transduction, transcription and effector functional gene categories as defined in Bettencourt et al. [50].
Of the four challenging conditions V. parahaemolyticus infections resulted in the highest number of genes with higher level of expression during this comparison study based on qPCR and selected genes targeting immune recognition, signal transduction, transcription, and synthesis of effector molecules processes (Figure 5). Also, cross‐talk between signaling pathways may occur in B. azoricus individuals subjected to Vibrio infections, wound responses, and hyperbaric stimulations, i.e., same immune or pro‐inflammatory signaling molecules may serve different signaling pathways whether they are conspicuously more expressed or not during such experiments. Cleary, the activation of signaling pathways involved in Vibrio infections was distinct from that of wound and hyperbaric reactions and thus conferring the animal model presented here with the physiological versatility to cope with deep‐sea hydrothermal vent environments. These experiments were important to elucidate the molecular mechanisms under which, physiological responses to bacterial infections, would responses, hyperbaric stimulations and long‐term maintenance in aquaria conditions, may be involved in B. azoricus adaptation processes whether in deep‐sea vent environments or at atmospheric pressure. However, in‐depth analysis of different signaling genes and pathways involved in such experimental challenges remained fragmentary and elusive.
\nOne the most common goals of RNA Sequencing (RNA‐Seq) profiling is to identify genes or molecular pathways that are differentially expressed (DE) between two or more biological conditions [59–63]. Changes in expression can then be associated with differences in physiological reactions, providing clues for further investigation into potential mechanisms of action [64, 65]. In order to gain additional insight into the different signaling genes involved in Vibrio infection, wound response, long‐term acclimatization, and hyperbaric repressurization, we sequenced the full transcriptome of gill tissues from each of these experimental challenges to which deep‐sea vent mussels were subjected and compared their differential gene expression levels with that of gene expression in animals immediately retrieved from the vent sites with the help of acoustically trigged cages that were recovered at the sea surface. Transcript sequences for the five cDNA libraries were obtained from the Illumina RNA‐sequencing platform and de novo assembly of RNA‐Seq transcripts performed with Trinity [66, 67] followed by differential expression (DE) analyses using the edgeR package [68–70]. DE results were presented as Heatmaps clusters (transcriptional cluster report for edgeR DE analysis). The advantage of Heatmaps is that it can display the expression pattern of the genes across all the RNA samples. Visualization of the results is aided by clustering together genes that have correlated expression patterns [68].
\nHere we present examples of expression plots for some of the most DE genes across the five different experimental conditions referred to as “cage,” animals freshly collected with acoustically triggered cages, from the bottom of the deep‐sea vent floor; “3 months,” same animals as in “cage” acclimatized for 3 months in aquaria environment at 1 atm; “Vibrio,” same animals as in 3 months exposed to V. parahaemolyticus; “Wound,” same animals as in 3 months with shell injury caused by mechanical abrasion to expose the mantle; “IPOCAMP,” same animals as in 3 months subjected to 80 bar hydrostatic pressure for 72 h. The top‐scoring BLAST hit for each of the gene exemplified is shown on top of the respective expression plot (Figure 6).
\nExpression plots across the five experimental conditions, “3 months”; “IPOCAMP”; “cage”; “Vibrio,” and “Wound,” representing differential gene expression analyses using EdgeR. The top‐scoring BLAST hit for each of the genes exemplified is shown on top of the respective expression plot.
Comparison of DE across the five experiment revealed interesting correlations as for “cage” and “3 months” mussels indicating that vent mussels endured well aquarium conditions for as long as 3 months, as demonstrated by similar levels of gene expression. Vibrio infections and IPOCAMP pressurization also showed clustering patterns of gene expression which would seem to indicate that once mussels are acclimatized to atmospheric pressure, repressurization stimulus is impacting vent mussels in similar ways as in V. parahaemolyticus challenges, suggesting thus the occurrence of stress‐related reactions in both types of stimulations. The expression pattern seen for wound injury was particularly distinct as compared to the other four experimental conditions. Wound injury seemed to affect drastically the vent mussel transcriptional activity which some of its genes were severely downregulated probably due to the damaging effect caused by the mechanical abrasion and direct exposure of the mantle to the aquarium environment. Taken together these experiments proven to be insightful in demonstrating the contrasting behavioral expression of given important physiological transcripts such as the peptidoglycan recognition and LBP‐BPI proteins, both involved in innate immune responses, when vent mussels are met with distinct environment factors.
\nThe transcriptome sequencing of gill tissues from the mussel B. azoricus revealed a set of genes of bacterial origin, providing a functional insight into the microbial vent community [71]. The transcripts supported a metabolically active microbiome and a variety of bacterial mechanisms and pathways, among which the fixation of carbon, the use of nitrate as a terminal acceptor of electrons and oxidation of sulfur and methane. The bacterial genes ensued from this sequencing work were deemed relevant to evaluate the influence of abiotic and biotic environmental conditions on B. azoricus transcriptional activity and also potentially useful to assess symbiont density differences in vent animals originated from distinct hydrothermal vent sites, respectively, to their environmental settings [72]. Keeping in line with the assumption that geographically distinct vent mussels will adopt different physiological statuses in relation to their environmental conditions, we also surmised that the relative abundance of methanotrophic and sulfide oxidizing endosymbiotic bacteria would differ between the Menez Gwen and Lucky Strike mussels as previously reported by other researchers [12, 58, 73]. We hypothesized that geographically distinct B. azoricus individuals may be experimentally traced back to their original hydrothermal vent sites based on their bacterial transcriptional activity and bacterial gill densities at the time animals were retrieved from the shallower Menez Gwen and deeper Lucky Strike vent sites. A taxonomical structure of the vent mussel gill’s microbiome was also assessed to determine the bacterial community composition of gill tissue from MG and LS mussels to infer the symbiont densities differences between animals from both vent sites. Results from the ribosomal RNA amplicon sequencing of the V6 hypervariable regions, by massive parallel 454 pyrosequencing, indicated that the percentage of sequences obtained was from endosymbiont bacteria at nearly the same proportion between Menez Gwen and Lucky Strike samples. Moreover, comparative analyses based on BLAST searches in the RDP database, using the 16S rRNA OTU sequences, revealed that the thiotrophic endosymbiont represented 90% of all the sequences and methanotrophic endosymbiont almost 5% of the sequences from vent mussel samples originated from the distinct Menez Gwen and Lucky Strike hydrothermal vent fields [72].
\nWhile the majority of our experiments using live vent mussels were performed shortly after their retrieval from the Menez Gwen hydrothermal vent, long‐term studies with vent mussels acclimatized to atmospheric pressure conditions have hardly been addressed until recently. As above‐mentioned, long‐term acclimatization experiments in aquarium systems have allowed us to study the expression of bacterial symbionts genes, particularly methanotrophic and thiotrophic bacteria, over time of acclimatization while their mussel host is faced with drastic physiological challenges, metabolic adaptations, and food intake changes in an effort to adapt to an aquarium environment at atmospheric pressure and without supplementation of methane and sulfur [56]. The physiological adaptation to aquarium environment is likely to be aggravated by the expelling of endosymbionts into the aquarium environment, progressively emptying the gill tissue of its autotrophic bacteria, essential for the host vent mussel nutritional sustenance. Long‐term aquarium acclimatization represents thus a model study to investigate the presence and maintenance of symbiotic associations between chemosynthetic bacteria and vent animals, which depend on controlled cell‐cell communication between host and endosymbionts and the role of the host immune system [56, 74].
\nPresumably, the loss of endosymbiont induces a dramatic change in host gene expression profiles especially if endosymbiont genes exert some transcriptional control over host gene expression. For this reason, acclimatization studies have been instrumental to further our understanding of B. azoricus immune system. These studies have provided insights into physiological principles underlying mechanisms of adaptation to aquarium conditions at sea level pressure while taking advantage of the remarkable capacity of vent mussels to survive well decompression once brought to surface [21, 22, 56]. Furthermore, these studies have allowed analyses using immune challenged mussels comparatively to acclimatized control mussels, maintained under aquarium conditions. In view of our previous experiments performed with live gill tissues and postcapture immune gene expression studies in B. azoricus acclimatized to atmospheric pressure, the presence of endosymbiont bacteria is now being under investigation as a driving factor under which host‐immune genes may transcriptionally be modulated and reciprocally endosymbiont genes may transcriptionally be modulated by the host [53–56]. Moreover, the impact of aquarium acclimatization on B. azoricus immune responses and its capacity to react to V. diabolicus challenges was recently evaluated during recurrent incubations with V. diabolicus during short periods of time, followed by clean sea water incubations allowing animals to depurate and subsequently be reexposed to the same load of V. diabolicus over a period of 3 weeks acclimatization experiment [74]. As previously described, we found a time‐dependent immune gene response in B. azoricus tied to the endosymbiont presence inside the vent mussel gills. The vent mussel’s immune defense capabilities were affected by the gradual loss of symbiont bacteria suggesting a symbiont‐mediated defense mechanism under which the transcriptional regulation of host immune genes is directly affected by symbiont density and/or activity. The host‐immune system‐endosymbiont interactions were actively higher during the first week of acclimatization as a result of Vibrio exposures, demonstrating the ability of B. azoricus to increase the transcription of immune genes while endosymbiont gene expression also correlated with an increased symbiotic metabolism and prevalence. A synergistic response was proposed to counteract the presence and potential infection by V. diabolicus bacterium while modulating B. azoricus immune defenses‐endosymbiont interactions to an extant, which host‐immune and endosymbiont genes are mutually reliant during the first weeks of acclimatization [74]. The evidence presented suggests successful V. diabolicus recognition prompting immune genes to increase their levels of transcriptional activity particularly for genes involved in the Toll‐like receptor signaling [75, 76] and apoptosis‐related pathways [77] during first day of acclimatization in aquarium environments. In agreement with this, B. azoricus is presented as a suitable model to study molecular interactions involving host‐mediated immune recognition events and adaptation mechanisms, to mitigate apoptosis harmful effects induced by Vibrio exposure against which, endosymbionts were prompted to increase their transcriptional activity, evocative of a possible protection role to the host [74]. This work brings to light other questions relating to how the host‐immune system regulates the symbiont population within their gills and conversely symbionts avoid being recognized and eliminated by the host. These topics are being further investigated in our group and focused on finding and characterizing the molecular mechanisms underlying the establishment (recognition and acquisition) and functioning of symbiosis between deep‐sea vent mussel B. azoricus and the methanotrophic and thiotrophic bacteria (gene expression, energy metabolism, regulation of symbiont population).
\nInterestingly, the study of intricate associations with chemosynthetic symbiont‐bacteria living in the gills of deep‐sea vent animals led us to the conception of a new pathogenesis model system based on an unconventional host‐symbiont model system. This new marine invertebrate model system, as for the ecotoxicological model Mytilus spp. [78] relies, instead, on its unique host‐immune‐symbiont bacteria interactions believed to play a crucial role in counteracting infectious pathogens. The establishment of invertebrate host pathogen systems may serve as suitable and useful models to study pathogenicity. The molecular mechanisms through which pathogens are able to colonize and overtake host’s immune system, particularly during the initial phase of infection when molecular recognition of MAMPs is occurring, as the pathogen defines its route of entry, are expected to reveal new molecular strategies that could help developing new therapies in aquaculture diseases. Using the deep‐sea vent mussel B. azoricus as an alternate invertebrate model system to study pathogenesis brings a new perspective into the search for new drug targets that could directly interfere with pathogen recognition processes and/or with in situ inflammatory process where immune cells (i.e., hemocytes) and cytokine‐like molecules are being mobilized. Indeed, in such host‐endosymbiont model systems, the role of endosymbiont‐derived molecules could have an important influence in mediating pathogenesis and in counteracting the deleterious effect of pathogens on the host immune system. From an experimental approach, several genera of bacterial fish pathogens may be used in B. azoricus, as infectious agents, e.g., Vibrio, Flavobacterium, Pseudomonas, Aeromonas, Streptococcus. Host and endosymbiont gene expression profiles may be studied during infection experiments carried with a given bacteria and genes that are markedly upregulated or downregulated further analyzed and their cDNA sequences determined by traditional sequencing methods.
\nParticular attention should be given to genes whose encoded proteins are participating in signal transduction pathways directly influencing the outcome of immune effector molecules, as antibacterial peptides; immune recognition lectins and antioxidant products such as superoxide dismutase, ferritin, metalloproteinases, metallothioneins, and heat shock proteins [26]. Synergistic effects resulting from interactions between host immune and endosymbiont activity, in counteracting infectious pathogens, may now be studied at the molecular level, for future therapies design, targeting key steps during pathogen infection processes, for instance, host recognition events; production of the anti‐inflammatory factor TNF‐alpha and cytokine‐like growth factors; enhancement of antibacterial molecules synthesis.
\nIn an attempt to understand physiological reactions of animals normally set to endure extreme conditions, in deep‐sea environments, our laboratory has undertaken, for the last 6 years, a series of investigations aimed at characterizing molecular indicators of adaptation processes of which components of the immune and antioxidative stress response systems have received most of our attention. As a research goal, long‐term maintenance of vent mussels to atmospheric pressure was instrumental to further our understanding on molecular relationships under which the vent mussel‐endosymbiont interactions are affected by aquaria conditions and by the gradual disappearance of endosymbiont bacteria from gill epithelia. Hence, the maintenance of live mussels in our aquarium laboratory system has been a key factor in gaining knowledge into the physiology of vent animals including the study of evolutionary conserved immune, inflammatory, and stress‐related factors commonly found in other marine bivalves. In vivo and ex vivo experiments conducted with live mussels and their excised gill tissues as primary tissue cultures, allowed the specific host‐endosymbiont interactions to be revealed, and further characterized in the deep‐sea vent model B. azoricus, establishing distinct genetic signatures for the expression of endosymbiont genes and host‐immune genes in relation to different environmental conditions. Increasing evidence now support the role of gills as a bone fide immune‐responsive tissue in B. azoricus, consistent with a suitable study model for exploring molecular interactions involving host‐endosymbiont‐mediated immune recognition events and adaptation mechanisms to deep‐sea hydrothermal vent environments. Such adaptation mechanisms are likely to be influenced by the microbial community composition surrounding the mussel beds at hydrothermal vents and therefore it is important to continue metatranscriptomic and metagenomic studies [79] from the gill‐associated microbial diversity and surrounding hydrothermal vent sediments [80, 81] in view of the broader ecological organization and evolutionary importance of animal‐bacterial microbiomes in chemosynthetic‐based ecosystems in the deep sea [82, 83].
\nIn recent years, researchers have turned to the human microbiome for its functional role in human health [84] and both composition and alterations in the microbiome have been found associated with diabetes, inflammatory bowel disease, obesity, asthma, rheumatoid arthritis, and susceptibility to infections [85]. Other microbiomes from nonmammalian and nonvertebrate species have also been characterized, for instance in insects where it was found to be highly dependent on the environment, species, and populations and affecting the fitness of species. These fitness effects may have important implications for the conservation and management of species and populations [82, 83]. Given the temporal instability of deep‐sea hydrothermal vents and their constant fluctuations of physical and chemical environmental conditions, vent animal‐microbiome associations have become critical for our understanding of invasion of nonnative species, responses to pathogens, and responses to chemicals and global climate change in the present and future [82] particularly when deep‐sea mining activities are projected to have a major impact on deep‐sea vent ecosystems [86].
\nAcknowledgements are due to IMAR—Marine Institute, Research and Development Unit #531 in the Azores (Thematic Area E) and Portuguese Fundação para a Ciência e Tecnologia (FCT) through the funding program FCOMP‐01‐0124‐FEDER‐007376 and the pluri‐annual and programmatic funding schemes—OE‐FEDER‐POCI2001‐FSE and PEst project—OE/EEI/LA0009/2011–2014. We also acknowledge the Regional Government of the Azores and Its Science and Technology Directorate through the pluri‐annual funding scheme and its funding support through the Operational Program PROCONVERGENCIA 2007‐20013 under the auspices of the European Union Regional strategic fund FEDER and through the research grant IBBA M2.1.2/I/029/2008‐BioDeepSea awarded to RB. The Luso‐American Foundation FLAD research grants project 600‐08/2016 and project L‐V‐173/2006; the Fulbright Commission through the Fulbright Research Scholar grant awarded to RB (2005‐2006) are also acknowledged. This work also received the support from the Fundação para a Ciência e Tecnologia, through the strategic research fund UID/MAR/04292/2013 granted to MARE‐Marine and Environmental Sciences Centre; through the research grant IMUNOVENT PTDC/MAR/65991/2006 awarded to RB; through the research grants DEEPFUN‐PTDC/MAR/111749/2009 and FCOMP‐OE/EEI/LA0009/2011) awarded to AC; through the post‐doctoral scholarship SFRH/BDP/14896/2004 awarded to RB; through the post‐doctoral scholarship SFRH/BPD/73481/2010 awarded to IM; through the doctoral scholarships SFRH/BD/689511/2010, and EM SFRH/BD/73152/2010 awarded to EM and IB, respectively. The Azorean Science and Technology Directorate doctoral scholarship M3.1.2/F/052/2011 granted to TC is acknowledge. RB was also supported by Fundação para a Ciência e Tecnologia through CIÊNCIA‐2007 Research contract (2008‐2013). AC was supported by the Program Investigador FCT‐IF/00029/2014/CP1230/CT0002 from the Fundação para a Ciência e Tecnologia.
\nThe taxing impact the fashion industry has had on the environment is by no means a new revelation—having accumulated a great deal of evidence over the years. However, unlike in the past when “sustainability” seemed more like an ideal adopted by individual, niche grassroot organizations, it is now considered a core value globally across the fashion industry. The fashion industry’s recent wave of intentional action toward sustainability is in part motivated by several comprehensive and revealing industry sustainability reports released in the last 3 years [1, 2, 3], but moreover it is a collective response to the recent fashion industry-specific sustainability campaigns such as the “2020 Commitment,” spearheaded in the last 2 years by several sustainability-driven coalitions (e.g., the Global Fashion Agenda and the Waste and Resources Action Programme UK), which have rallied formal commitments from a significant portion of the fashion industry toward concrete, quantifiable action for sustainability by 2020.
The heightened concern toward the fashion industry’s environmental impact is also stirred by evidence of intensified global clothing consumption—which according to data from the World Bank [4] has doubled from around 50 billion units of clothing sales in 2000 to over 100 billion units in 2015 (see Figure 1). This dramatic increase in clothing consumption has been fueled by fast fashion, an increasingly bargain-driven consumer, increased accessibility via an expanding online shopping landscape, and more buying power from a growing middle class, especially in emerging economies such as China (projected to surpass the United States “as the largest fashion market in the world” in 2019, according to McKinsey FashionScope [5]). Unfortunately, the increased accessibility and affordability of clothing simultaneously propagated not only a culture of excessive consumption but also a quicker disposal of clothing, as exemplified by an approximately 20% decrease in the average number of times a garment is worn before it is abandoned as shown in Figure 1.
Growth of clothing sales and decline in clothing utilization since 2000. Source: World bank, [4].
Given the abundance of information surrounding the subject of sustainability in the fashion industry from many sources, there is an opportunity for a collated overview on the subject. Therefore, the purpose of this article is to provide an overview of (1) the most concerning environmental impacts caused by the fashion industry, (2) current leading collective sustainability campaigns mobilizing the fashion industry, (3) current available benchmarks and tools for measuring environmental impact of the textile life cycle, and (4) examples of how companies in the fashion industry are executing sustainability initiatives in their products or processes. Finally, the article will conclude with some of the current challenges and future opportunities in sustainability confronting the fashion industry.
In any given industry, each stage of the product life cycle poses an impact on the environment—by consuming environmental inputs (e.g., water for harvesting raw materials, fossil fuels to power manufacturing equipment, etc.) and releasing environmental outputs (e.g., carbon dioxide emissions from burning fossil fuels, landfill waste after product is disposed, etc.). For the fashion industry, the environmental inputs and outputs of the textile product life cycle is reflected in Figure 2. (It is worthwhile to note that the term “life cycle” used is misleading in that the above chain of processes does not form a “cycle,” but is instead linear sequence of events, with a definite beginning and end. A true cyclical life cycle would be indicative of recycling or reuse, feeding the end waste back into the system to be used again).
Environmental impact (inputs and outputs) of the textile life cycle.
As shown, the inputs and outputs of the fashion industry’s “textile product life cycle” pose impact on the environment, but it is the size of the impact which is staggering. This is partly due to the immense scale of the fashion industry, which has been evaluated to be a USD 1.3 trillion dollar industry [6], and the world’s third largest manufacturing industry, after automotive and technology [7]. But also, according to a report by the Ellen MacArthur Foundation, data confirms that the greenhouse gas emissions produced by textile production exceeds that of international aviation and maritime shipping combined. If it continues down this path, it is projected that by 2050 it could account for 1/4 of the worlds’ carbon emissions [1]. To put it into perspective further, the annual carbon footprint of the fashion industry’s product life cycle (3.3 billion tons CO2 emissions) is almost equivalent to that of 28 countries in the EU (3.5 billion tons) [7].
However, greenhouse gas emissions are not the only harmful environmental outputs from the fashion industry; it is just one of the numerous other inputs and outputs which have strenuous environmental implications, as exemplified in Figure 2. The below provides a summary, along with examples, highlighting some of the leading concerns (note that there are indeed many others; however, for the purpose of this condensed article, we will focus on the following):
Heavy consumption of depleting natural resources:
For example, water consumption for cotton crops
For example, coal/natural gas (nonrenewable) energy to power manufacturing facilities
Polluting waste outputs (e.g., chemicals, pesticides, carbon emissions, etc.):
For example, fertilizer/pesticide runoff from cotton crops
Dyes/chemical waste from garment factories (e.g., for dyeing and washing processes)
Microplastic pollution (e.g., from synthetic fiber shedding):
For example, shedding of polyester fibers (considered microplastics) in the laundry process: a domestic wash load can release around 700,000 fibers and, as they are unable to be completely filtered out by waste water treatment plants, end up infiltrating and accumulating in marine ecosystems [8]. This issue is exacerbated by the drastic increase in the annual consumption of polyester fibers in the fashion industry, which has grown exponentially, from 8.3 million tons in 2000, to 21.3 million tons in 2016 [6].
This section provided a condensed overview of the extent of the fashion industry’s impact on the environment and highlighted the most concerning forms of impact. However, it is worth noting that the abundance of published data and literature on the environmental impact of the fashion industry is truly inundating and could easily extend beyond the scope of this section. The following section will present some of the current collective global sustainability campaigns which are striving to alleviate the environmental impact of the fashion industry in the future.
The intensified evidence of the fashion industry’s impact on the environment in the last decade prompted the founding of several global sustainability campaigns within the last 3 years. These campaigns, spearheaded by sustainability-driven coalitions, are mobilizing companies across the fashion industry, collectively toward adopting sustainable materials and practices throughout their design, development, and supply chains, and have already garnered formal commitments from key players in the fashion industry which represent a sizable portion of the market. Two predominant global campaigns, initiated in 2018, are summarized below:
The “2020 Circular Fashion System Commitment,” introduced by the Global Fashion Agenda
Mission/action points:
The Global Fashion Agenda is a leadership forum engaging the fashion industry toward sustainability [9]. Its “2020 Circular Fashion System Commitment” is a call on the fashion industry to commit toward a “circular fashion system,” by taking concrete action on one or more of the following points:
Implementing design strategies for cyclability
Increasing the volume of used garments and footwear collected
Increasing the volume of used garments and footwear resold
Increasing the share of garments and footwear made from recycled post-consumer textile fibers
Industry commitment (as of May 2018):
Ninety-four companies signed on (represents 12.5% of the global fashion market), including ASOS, H&M, Nike, Inditex, Kering, and Target.
The “Sustainable Clothing Action Plan (SCAP) 2020 Commitment,” introduced by the Waste and Resources Action Programme (WRAP)
Mission/action points:
The SCAP (spearheaded by WRAP) is a collaborative framework and voluntary commitment for organizations to deliver industry-led targets of a 15% reduction in carbon, water, and waste in the clothing industry by [10]:
Reinventing how clothes are designed and produced
Rethinking how we value clothing by extending life of clothes
Redefining what is possible through reuse and recycling
Industry commitment (as of March 2019):
Eighty companies signed on (represents 58.5% of the UK’s retail sales volume), including ASOS, Marks and Spencer, Ted Baker, and others.
The action points of both these campaigns show an emphasis on cyclability—not just of materials but also practices—and reshaping the product life cycle toward circularity [10] (see Figure 3). The number of companies committed to these campaigns so far is a promising sign that sustainability is gradually becoming an integral factor in the fashion industry. Aside from the global sustainability campaigns such as above, another industry resource supporting companies toward sustainability is the various benchmarks and tools developed to help the fashion industry gauge the environmental impact of certain materials or processes and therefore help steer decisions accordingly. The following section will explore some of these tools and benchmarks.
For companies in the fashion industry to become more cognizant and proactive about minimizing the environmental impact of their product life cycles, they would need to rely on a definitive benchmarks and tools to gauge the environmental impact of their decisions regarding product or processes. However, measuring environmental impact of such decisions can be very convoluted, as results tend to be conflicting depending on which angle it is viewed from. Here are some examples of the conflicting nature of environmental impact measures:
On the one hand, for example:
A polyester shirt has more than double the carbon footprint of a cotton shirt (5.5 kg CO2 emissions vs. 2.1 kg CO2 emissions) [11].
But on the other hand:
The processing for cotton produces a water footprint 20 times larger than that of polyester (see Figure 4).
One kilogram of cotton—equivalent to the weight of a shirt and pair of jeans—can take as much as 10,000–20,000 liters of water to produce [10].
For an organic cotton tote to make up for the environmental impact (water use, energy use, etc.) of a classic plastic bag, it would need to be used 20,000 times [12].
A diagrammatic expression of the goal of “circularity” in the textile product life cycle.
The following is an outline of three established benchmarks and tools, designed to enable the fashion industry (and other industries), to measure the environmental impact of certain decisions regarding their material use or processes employed:
Higg Index, developed by the Sustainable Apparel Coalition:
It is described as “a suite of tools” that enables the measure and score of a company or product’s “sustainability performance” at “every stage in their sustainability journey,” aiming to provide a “holistic overview” that “empowers businesses to make meaningful improvements that protect the well-being of factory workers, local communities, and the environment” [13]. It encompasses the following tools:
Product tools:
Higg Materials Sustainability Index (MSI): “the apparel industry’s most trusted tool to accurately measure the environmental sustainability impacts of materials,” by scoring materials based on their environmental impact from fiber to fabric across five environmental impact parameters (global warming, water pollution, water scarcity, resource depletion, and chemicals) (see Figure 5 for a sample screenshot of the Higg MSI interface)
Higg Design and Development Module (DDM): “guides designers to combine their chosen materials for maximum positive impact, to select the most sustainable manufacturing techniques, and to consider the complete life-cycle of the product”
Higg Product Module (PM): will measure the environmental impact a product (apparel, footwear, and textile products) generates throughout its life cycle when produced at industrial scale and be able to cross compare products with one another as well as which life cycle stages or production processes contribute the most impact (expected to launch in 2019)
Facility tools:
Higg Facility Environmental Module (FEM): measures the environmental impact of individual factories based on assessing factors such as environmental management systems, energy use and greenhouse gas emissions, water use, wastewater, emissions to air (if applicable), waste management, and chemical use and management
Higg Facility Social and Labor Module (FSLM): measures the social impact of individual factories based on assessing factors such as recruitment and hiring, working hours, wages and benefits, employee treatment, employee involvement, health and safety, termination, management systems, facility workforce standards and those of value chain partners, external engagement on social and labor issues with other facilities or organizations, and community engagement
Brand and retail tools:
Higg Brand and Retail Module (BRM): enables brands and retailers of all sizes to measure the environmental and social and labor impacts of their operations across a product’s life cycle (from materials sourcing through its end of use). The environmental impacts measured include greenhouse gas (GHG) emissions, energy use, water use, water pollution, deforestation, hazardous chemicals, and animal welfare. The social and labor impacts measured include child labor, discrimination, forced labor, sexual harassment and gender-based violence in the workplace, non-compliance with minimum wage laws, bribery and corruption, working time, occupational health and safety, and responsible sourcing.
MADE-BY Environmental Benchmark for Fibers, developed by MADE-BY in cooperation with Brown and Wilmanns Environmental, LLC:
It ranks 28th in the most commonly used fibers in the garment industry into 5 classes (Class A–E), based on the following measures: greenhouse gas emissions, human toxicity, eco-toxicity, energy, water, and land [15] (see Figure 6).
Corporate Fiber and Materials Benchmark (CFMB) (formerly the Preferred Fiber and Materials Benchmark (PFMB)), launched by the Textile Exchange:
Launched in 2015, it is a leading industry-led, voluntary self-assessment tool which enables companies to systematically measure, manage, and integrate a preferred fiber and materials strategy into four key areas of mainstream business operations: corporate strategy, supply chain, consumption, and consumer engagement [16] (see Figure 7 for flowchart of this framework laid out). It also provides feedback on progress and performance in comparison to peers and the overall industry. As of 2018, 111 companies have partaken in the program (an increase of 106% since 2015).
Water footprint for the total processing phase of each fiber type for the UK (m3) in 2016. Source: Waste and resources action programme [14].
Sample screenshots of the Higg Materials Sustainability Index (MSI) tool. Source: Sustainable apparel coalition [13].
The MADE-BY environmental benchmark fiber classification chart. Source: Common objective [15].
As can be seen from the three examples above, there is a wide selection of benchmarks and tools for measuring environmental impact available to the fashion industry; however, there are some limitations to consider. For one, the wide selection can also be problematic as each of the different initiatives above accounts for slightly different factors or weighs them slightly differently; therefore the result obtained from one tool might not be consistent with that obtained from another. For example, based on the Higg Materials Sustainability Index, natural fibers like silk, cotton, and wool are assigned higher environmental impact scores (i.e., more damaging to environment) of 128, 98, and 82, respectively, while fossil-fuel-derived fibers like nylon, acrylic, and polyester have lower impact scores at 60, 52, and 44 [7]. This is because the Higg Index puts greater emphasis on fiber production, which is indeed more taxing on the environmental for natural fibers such as silk, cotton, and wool, as their procurement imposes a greater strain on natural resources (such as water, land, or animal welfare). Yet, in contrast, according to the MADE-BY Environmental Benchmark (Figure 6), fossil-fuel-based virgin nylon fibers and natural wool fibers are both ranked under the same Class E (the “least sustainable” category). Hence the availability of multiple benchmarks and tools could prove to be more incumbering than helpful when it comes to definitively measuring environmental impact.
Another limitation of these benchmarks and tools is that they do not sufficiently weigh in, or even overlook, the impact of the in-use phase of the textile product life cycle. The in-use phase here refers to the period when the textile product is being used for what it was made for. So, for a garment, that would mean the period from when it is purchased by a customer until it is no longer used or disposed of, which mostly involves its wearing and laundering. The research of Laitala et al. reveals that energy and water consumption during the laundering process varies greatly depending on fiber content of the garments [17]. Firstly, (see Figure 8) the research presents data which indicates that wool- and silk-based garments are 3–6 times more likely to be dry-cleaned than cotton- or synthetic-based garments and furthermore that dry cleaning uses 3–6 times (depending on the type of dry-cleaning process) more electricity than wet washing methods (which is the predominant laundering method for cotton- or synthetic-based garments). However, their research also shows that on average, the water temperature of the wash setting for cotton-based garments is about 17°C higher than that for wool-based garments. With polyester and other nonbiodegradable polymer fibers (e.g., acrylic and nylon), there is the developing concern regarding the shedding of fibers (microplastic) during the washing process which, being unable to be completely filtered out by standard waste water treatment plants, end up infiltrating and accumulating in marine ecosystems.
Flowchart showing the framework of the textile exchange’s corporate fiber and materials benchmark (CFMB). Source: Textile exchange, [16].
Another aspect which deserves more consideration by the benchmarks and tools is human ecology and not just environmental ecology. For example, there are man-made fibers derived from plants, such as polylactic acid (PLA) derived from corn, which are environmentally biodegradable, but not necessarily human biocompatible [18]. Therefore, the potential negative side effects or toxicity on human ecology is a factor which deserves equal attention in impact measures.
These limitations in the current benchmarks and tools are a clear reminder that measuring environmental impact of product or processes in the fashion industry is multifaceted and convoluted. Currently there is no prevailing, overriding benchmark or tool that provides a definite unanimous measure of environmental impact, so it is up to companies to adopt a holistic approach when developing a strategy toward sustainability.
Having reviewed several sustainability campaigns and environmental impact measure benchmarks and tools relevant to the fashion industry today, this section will now proceed to provide insight into how companies and various players in the industry have responded, i.e., the kinds of strategic initiatives being taken toward sustainability. The sustainability initiatives will be categorized into two types: (1) front-end approach and (2) back-end approach.
Within the context of this article, this refers to the integration of sustainable initiatives at the beginning stages (front-end) of the textile product life cycle, such as in the raw material sourcing and design and development processes. So, for example, a front-end sustainable initiative could be the decision to use “low environmental impact*” textile fibers as the raw materials for the textile goods being produced. A front-end sustainable initiative could also be manifested in the design and development process, for example, by utilizing digital tools to minimize the need for physical prototype samples or by training designers to adopt an eco-conscious mindset into their creations. (*Note that we are using the term “low environmental impact” textile fibers as opposed to “sustainable” or “eco-friendly” or “green” textile fibers because the latter terms can be misleading as there are no completely “sustainable/eco-friendly/green” fibers; all materials pose some impact. Furthermore, as discussed in the previous section, it is difficult to resolutely confirm the impact of a certain material, as there are many facets of environmental impact. Therefore “low environment impact” is a more accurate representation of what is possible to strive for in sustainable materials).
An industry example of a front-end approach to sustainability is the adoption of regenerated cellulosic fibers, such as Lyocell and Seacell, by various fashion companies particularly in lingerie and activewear [19]. With cotton, albeit a natural cellulosic fiber, bearing a hefty water footprint in the harvesting process, and with petrochemical-based synthetic fibers such as polyester and nylon bearing a hefty carbon footprint in the manufacturing process [20], regenerated cellulosic fibers can prove advantageous. They have the benefit of being biodegradable and derived from natural renewable resources (i.e., Lyocell is derived from wood pulp and Seacell is derived from seaweed) via a closed-loop manufacturing process, thereby consuming far less water and energy than traditional cotton, polyester, and nylon. Both Lyocell and Seacell also naturally carry antibacterial and fast-drying properties, which is why they are ideal for lingerie and activewear product.
A limitation of a front-end approach in tackling environmental impact is that it is still feeding more product in the fashion pipeline which will eventually end up at the end of the textile life cycle as waste by-product (even if it is biodegradable by-product) which needs to be managed accordingly. Therefore, in the following section, we will look at an approach which tackles the by-product end of the textile product life cycle.
Within the context of this article, this is referring to sustainability initiatives which aim to minimize the environmental impact of the product and processes at the end of the textile product life cycle, e.g., at disposal. A prime example of this is exemplified in the now widespread initiatives of post-consumer textile recycling. The reason recycled textiles have become so prevalent as a strategy to minimize environmental impact is not only because of the exponential supply of textile waste driven by intensified clothing consumption but more strategically because research has shown that the fiber production stage (extraction and processing) of the textile product life cycle has the greatest environmental impact in terms of water and carbon footprint, as shown in Figures 9–10 [14]. Therefore, by recycling post-consumer textile waste back into the textile supply chain enables bypassing the heavy environmental toll of the fiber production stage.
Data on laundry requirements based on fiber content. Source: Laitala et al. [17].
Water footprint of clothing in the UK (m3) in 2012 and 2016, comparing life cycle stages. Source: Waste and resources action programme [14].
Carbon footprint of clothing in the UK (t CO2e) in 2016, by process. Source: Waste and resources action programme [14].
There has been a great deal of research invested into textile recycling, from both the industry and academia. One notable advancement in textile recycling is exemplified by Garment-to-Garment (G2G) Recycle System, a closed-loop garment recycling retail concept supported by technology which enables the recycling of blended post-consumer garments, developed by HKRITA, in partnership with H&M and Novetex [21]. The Garment-to-Garment (G2G) Recycle System brings garment recycling to the retail level, therefore paving the way for garment recycling to be more accessible to the everyday consumer.
There are also several notable recycling initiatives which, instead of relying solely on post-consumer textile products, are derived from various kinds of post-consumer plastic waste. REPREVE is one example of this. Produced by the company Unifi, REPREVE is a brand of polyester fibers made from recycled post-consumer plastic waste (e.g., plastic bottles) [22]. The ability to convert various forms of plastic waste into usable polyester textile fibers has the benefit of resourcefulness. Even though the conversion from post-consumer plastic waste to fiber requires energy and water input for the manufacturing process, according to Unifi, it is reportedly much less than that required for virgin polyester (energy consumption is 45% less, water consumption is almost 20% less, and over 30% less greenhouse gas emissions).
Over the past decade, there have been many encouraging advancements which have expanded back-end approach sustainability initiatives such as textile recycling. However, there remain limitations in the current textile recycling technologies. For example, due to the need for comprehensive shredding in breaking down post-consumer textile waste, the tensile strength of recycled cotton yarns is less than that of virgin cotton [23]. Furthermore, as recycled yarns are composed of a mixture of fibers which may have undergone different dyeing and finishing processes in their last life, even after cleaning and bleaching processes, they may not be able to achieve the same hand-feel and color vibrancy possible with virgin fibers, therefore limiting its design versatility. These are some examples of limitations which could be preventing a greater adoption of textile recycling in the industry.
This article has attempted to provide a current and overarching view on the most concerning environmental impacts of the fashion industry today, the leading global sustainability campaigns and benchmarks and tools established to help empower the fashion industry toward concrete action and, last but not least, examples of sustainability initiatives being implemented in the industry. The fashion industry’s large-scale wave of movement toward sustainability is evident; however, there remain questions and challenges to be addressed, one being how successful the “2020 commitment” goals will be, with 2020 just around the corner, and considering how potentially disruptive any kind of change is in an industry which is built on long-established processes and practices and adheres to an inflexible, tight calendar. Furthermore, as discussed in this article, the array of benchmarks and tools available for measuring environmental impact can result in a convoluted process and conflicting, inconclusive information. Such challenges may deter a company from successfully achieving concrete changes toward sustainability.
Even if companies are able to navigate through the intricacies in evaluating environmental impact of a textile product or process, it is important to remember that the textile product life cycle is never impact-free (at least not in the foreseeable future), as it relies on the environment to provide various inputs and outputs. With this reality in mind, companies may find that making small but carefully holistically considered steps in the right direction can be much more effective than larger uninformed leaps when it comes to sustainability.
This work was funded by the Hong Kong Research Institute of Textiles and Apparel Limited (HKRITA), grant ITP/001/18TP. We would also like to thank the team at Tory Burch LLC for sharing their insight and ideas regarding existing industry sustainability initiatives, which helped inform some of the references in this work (but are not accountable for any potential errors found in this work).
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