Sample outline of an evaluation plan with Kirkpatricks’ four levels.
\\n\\n
Released this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\\n\\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
Note: Edited in March 2021
\\n"}]',published:!0,mainMedia:{caption:"Highly Cited",originalUrl:"/media/original/117"}},components:[{type:"htmlEditorComponent",content:'IntechOpen is proud to announce that 191 of our authors have made the Clarivate™ Highly Cited Researchers List for 2020, ranking them among the top 1% most-cited.
\n\nThroughout the years, the list has named a total of 261 IntechOpen authors as Highly Cited. Of those researchers, 69 have been featured on the list multiple times.
\n\n\n\nReleased this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\n\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
Note: Edited in March 2021
\n'}],latestNews:[{slug:"webinar-introduction-to-open-science-wednesday-18-may-1-pm-cest-20220518",title:"Webinar: Introduction to Open Science | Wednesday 18 May, 1 PM CEST"},{slug:"step-in-the-right-direction-intechopen-launches-a-portfolio-of-open-science-journals-20220414",title:"Step in the Right Direction: IntechOpen Launches a Portfolio of Open Science Journals"},{slug:"let-s-meet-at-london-book-fair-5-7-april-2022-olympia-london-20220321",title:"Let’s meet at London Book Fair, 5-7 April 2022, Olympia London"},{slug:"50-books-published-as-part-of-intechopen-and-knowledge-unlatched-ku-collaboration-20220316",title:"50 Books published as part of IntechOpen and Knowledge Unlatched (KU) Collaboration"},{slug:"intechopen-joins-the-united-nations-sustainable-development-goals-publishers-compact-20221702",title:"IntechOpen joins the United Nations Sustainable Development Goals Publishers Compact"},{slug:"intechopen-signs-exclusive-representation-agreement-with-lsr-libros-servicios-y-representaciones-s-a-de-c-v-20211123",title:"IntechOpen Signs Exclusive Representation Agreement with LSR Libros Servicios y Representaciones S.A. de C.V"},{slug:"intechopen-expands-partnership-with-research4life-20211110",title:"IntechOpen Expands Partnership with Research4Life"},{slug:"introducing-intechopen-book-series-a-new-publishing-format-for-oa-books-20210915",title:"Introducing IntechOpen Book Series - A New Publishing Format for OA Books"}]},book:{item:{type:"book",id:"6313",leadTitle:null,fullTitle:"Neoplasm",title:"Neoplasm",subtitle:null,reviewType:"peer-reviewed",abstract:"Neoplasm refers to an abnormal tissue growth that arises as a consequence of rapid cell proliferation and continues to grow abnormally even after terminating the stimuli that had instigated the change. It lacks partial or complete functional coordination with that of the normal tissues. Neoplasms are classified depending on the degree and type of tissues involved. Carcinogenesis is a multistep process where a plethora of endogenous and exogenous factors turns out genetic and epigenetic modifications, which collectively amend some critical cellular pathways controlling the proliferation, apoptosis, and differentiation. The cells having aberrant modifications are transformed into malignant ones of which the clonal expansion results in the development of cancer. This book provides the reader with a comprehensive overview of various cancer types along with their molecular mechanisms of initiation and progression. It also describes the current knowledge about the state-of-the-art measures being employed in cancer diagnosis and therapeutics. Particular attention is paid to make this book equally useful for students, practitioners, and expert scientists.",isbn:"978-1-78923-778-8",printIsbn:"978-1-78923-777-1",pdfIsbn:"978-1-83881-393-2",doi:"10.5772/intechopen.69560",price:119,priceEur:129,priceUsd:155,slug:"neoplasm",numberOfPages:254,isOpenForSubmission:!1,isInWos:null,isInBkci:!1,hash:"dfea745f8ae5593e6dfc35d9e621291f",bookSignature:"Hafiz Naveed Shahzad",publishedDate:"September 19th 2018",coverURL:"https://cdn.intechopen.com/books/images_new/6313.jpg",numberOfDownloads:12931,numberOfWosCitations:33,numberOfCrossrefCitations:38,numberOfCrossrefCitationsByBook:0,numberOfDimensionsCitations:67,numberOfDimensionsCitationsByBook:0,hasAltmetrics:0,numberOfTotalCitations:138,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"June 12th 2017",dateEndSecondStepPublish:"July 3rd 2017",dateEndThirdStepPublish:"November 23rd 2017",dateEndFourthStepPublish:"December 28th 2017",dateEndFifthStepPublish:"February 26th 2018",currentStepOfPublishingProcess:5,indexedIn:"1,2,3,4,5,6",editedByType:"Edited by",kuFlag:!1,featuredMarkup:null,editors:[{id:"180702",title:"Dr.",name:"Hafiz",middleName:"Naveed",surname:"Shahzad",slug:"hafiz-shahzad",fullName:"Hafiz Shahzad",profilePictureURL:"https://mts.intechopen.com/storage/users/180702/images/5522_n.jpg",biography:"Dr. Hafiz Naveed Shahzad completed his PhD in 2013 from the International Agency for Research on Cancer (IARC), World Health Organization (WHO), France. Currently, he is working as an assistant professor at the School of Biological Sciences, University of the Punjab, Lahore, Pakistan. He is conducting research on multiple aspects of cancer and oncogenic viruses. His research expertise is in microbiology as well as cell and molecular biology. He has published more than a dozen research papers in high-impact factor journals and presented his work at various international conferences across the globe. Moreover, he has been serving as an editorial board member of various journals of international repute.",institutionString:null,position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"2",totalChapterViews:"0",totalEditedBooks:"1",institution:{name:"University of the Punjab",institutionURL:null,country:{name:"Pakistan"}}}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,coeditorOne:null,coeditorTwo:null,coeditorThree:null,coeditorFour:null,coeditorFive:null,topics:[{id:"428",title:"Cancer Biology",slug:"biochemistry-genetics-and-molecular-biology-oncology-cancer-biology"}],chapters:[{id:"61102",title:"Neoplasms of Central Nervous System: A Diagnostic Approach",doi:"10.5772/intechopen.76294",slug:"neoplasms-of-central-nervous-system-a-diagnostic-approach",totalDownloads:1276,totalCrossrefCites:1,totalDimensionsCites:1,hasAltmetrics:0,abstract:"Neoplasms of central nervous system accounts for approximately 1% of tumors of the human body, and they can be primary or secondary (metastatic), benign or malignant, and intra-axial or extra-axial. This chapter includes some most common brain and spinal cord tumors, like pituitary adenomas, meningiomas and gliomas, with their clinical, imaging, and histological characteristics for the diagnosis purpose, with additional treatment options and prognosis.",signatures:"Frank Yuan Shan, Dingrong Zhong, Wanming Hu, Nitesh Patel,\nEkokobe Fonkem, Dongxia Feng, Yilu Zhang, Jason H. Huang and\nArundhati Rao",downloadPdfUrl:"/chapter/pdf-download/61102",previewPdfUrl:"/chapter/pdf-preview/61102",authors:[{id:"224305",title:"Dr.",name:"Frank Y.",surname:"Shan",slug:"frank-y.-shan",fullName:"Frank Y. Shan"}],corrections:null},{id:"62306",title:"Uterine Sarcomas: An Updated Overview. Part 1: Smooth Muscle Tumors",doi:"10.5772/intechopen.76772",slug:"uterine-sarcomas-an-updated-overview-part-1-smooth-muscle-tumors",totalDownloads:1308,totalCrossrefCites:1,totalDimensionsCites:1,hasAltmetrics:0,abstract:"Uterine sarcomas (USs) account for 3–9% of uterine malignant neoplasia and about 5% of all gynaecologic malignancies. Despite their low prevalence, these tumors stimulate a great interest because of their aggressiveness, poor prognosis and high mortality rate. According to the last World Health Organization (WHO) classification and the International Federation of Gynecology and Obstetrics Committee (FIGO) staging, USs are categorized as pure mesenchymal tumors (endometrial stromal sarcoma, leiomyosarcoma and undifferentiated uterine), and mixed tumors (carcinosarcoma and adenosarcoma). Due to their non-specific signs and symptoms, USs are commonly diagnosed in advanced stage, more often after surgery for a suspected leiomyoma. Although surgery followed by adjuvant therapies represent the common choices for USs, they show poor efficacy due to the early occurrence of metastasis, and the high resistance of tumors to radio-and chemotherapy. Presently, specific expression profiles and new cytotoxic agents are under investigation. In these reviews, we summarized clinical and pathological features, imaging characteristics, therapeutic approaches, genomic and molecular aberration associated with smooth muscle neoplasia (Part 1) and endometrial stromal neoplasia (Part 2); the goal is to understand the biology and the molecular signature of these tumors, in order to focus on their best management.",signatures:"Roberta Zappacosta, Francesco Fanfani, Barbara Zappacosta,\nFrancesca Sablone, Lucia Pansa, Marco Liberati and Sandra Rosini",downloadPdfUrl:"/chapter/pdf-download/62306",previewPdfUrl:"/chapter/pdf-preview/62306",authors:[{id:"43833",title:"Prof.",name:"Sandra",surname:"Rosini",slug:"sandra-rosini",fullName:"Sandra Rosini"},{id:"80933",title:"Prof.",name:"Roberta",surname:"Zappacosta",slug:"roberta-zappacosta",fullName:"Roberta Zappacosta"},{id:"99169",title:"Dr.",name:"Barbara",surname:"Zappacosta",slug:"barbara-zappacosta",fullName:"Barbara Zappacosta"},{id:"218234",title:"Prof.",name:"Marco",surname:"Liberati",slug:"marco-liberati",fullName:"Marco Liberati"},{id:"218235",title:"Dr.",name:"Francesca",surname:"Sablone",slug:"francesca-sablone",fullName:"Francesca Sablone"},{id:"218236",title:"Dr.",name:"Lucia",surname:"Pansa",slug:"lucia-pansa",fullName:"Lucia Pansa"},{id:"218612",title:"Prof.",name:"Francesco",surname:"Fanfani",slug:"francesco-fanfani",fullName:"Francesco Fanfani"}],corrections:null},{id:"62297",title:"Uterine Sarcomas: An Updated Overview Part 2: Endometrial Stromal Tumor",doi:"10.5772/intechopen.78980",slug:"uterine-sarcomas-an-updated-overview-part-2-endometrial-stromal-tumor",totalDownloads:1213,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"Uterine sarcomas (USs) account for 3–9% of uterine malignant neoplasia and about 5% of all gynaecologic malignancies. Despite their low prevalence, these tumors stimulate a great interest because of their aggressiveness, poor prognosis and high mortality rate. According to the last world health organization (WHO) classification and the International Federation of gynecology and obstetrics committee (FIGO) staging, USs are categorized as pure mesenchymal tumors (endometrial stromal sarcoma, leiomyosarcoma and undifferentiated uterine) and mixed tumors (carcinosarcoma and adenosarcoma). Due to their non-specific signs and symptoms, USs are commonly diagnosed in advanced stage, more often after surgery for a suspected leiomyoma. Although surgery followed by adjuvant therapies represent the common choices for USs, they show poor efficacy due to the early occurrence of metastasis, and the high resistance of tumors to radio-and chemotherapy. Presently, specific expression profiles and new cytotoxic agents are under investigation. In these reviews, we summarized clinical and pathological features, imaging characteristics, therapeutic approaches, genomic and molecular aberration associated with smooth muscle neoplasia (Part 1) and endometrial stromal neoplasia (Part 2); the goal is to understand the biology and the molecular signature of these tumors, in order to focus on their best management.",signatures:"Roberta Zappacosta, Francesco Fanfani, Barbara Zappacosta,\nFrancesca Sablone, Lucia Pansa, Marco Liberati and Sandra Rosini",downloadPdfUrl:"/chapter/pdf-download/62297",previewPdfUrl:"/chapter/pdf-preview/62297",authors:[{id:"80933",title:"Prof.",name:"Roberta",surname:"Zappacosta",slug:"roberta-zappacosta",fullName:"Roberta Zappacosta"}],corrections:null},{id:"63256",title:"Novel Mechanism of Nonalcoholic Lipid Accumulation Promoting Malignant Transformation of Hepatocytes",doi:"10.5772/intechopen.77400",slug:"novel-mechanism-of-nonalcoholic-lipid-accumulation-promoting-malignant-transformation-of-hepatocytes",totalDownloads:1247,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"The incidence of hepatocellular carcinoma (HCC) is steadily increasing in worldwide, which has been a public concern significantly associated with diabetes and non-alcoholic fatty liver disease (NAFLD) is an emerging risk factor with increasing prevalence nowadays, with gradually instead of HBV and HCV, aflatoxin, or alcohol liver disease as major etiological factors. The deeply worrisome aspects of these high risk factors are their large spread in population. Systemic and genetic mechanisms involved in malignant transformation of liver cells as well as useful biomarkers at early stage of HCC are being investigated. However, the exact mechanisms from NAFLD to HCC still remain to be explored. In this paper, some advances of liver lipid accumulation were summarized on the relationship between NAFLD and hepatocytes malignant transformation.",signatures:"Min Yao, Wenjie Zheng, Li Wang, Miao Fang, Zhizhen Dong and\nDengfu Yao",downloadPdfUrl:"/chapter/pdf-download/63256",previewPdfUrl:"/chapter/pdf-preview/63256",authors:[{id:"32568",title:"Prof.",name:"Dengfu",surname:"Yao",slug:"dengfu-yao",fullName:"Dengfu Yao"}],corrections:null},{id:"60555",title:"Glucose Metabolism and Carcinogenesis: The Impact of the Tumor Suppressor p53",doi:"10.5772/intechopen.75976",slug:"glucose-metabolism-and-carcinogenesis-the-impact-of-the-tumor-suppressor-p53",totalDownloads:1108,totalCrossrefCites:2,totalDimensionsCites:2,hasAltmetrics:0,abstract:"The tumor suppressor protein, p53 responds to cellular stress such as DNA damage, oncogenic activation and hypoxia by transactivating downstream genes that are responsible for apoptosis, DNA repair, senescence, cell cycle arrest and cell cycle progression. However, emerging trends show that p53 also plays multifaceted roles in regulating glucose metabolism. It promotes oxidative phosphorylation, suppresses glycolysis at multiple points as well as controlling glutamine and lipid metabolism. Current findings suggest that p53 actions have potential to influence the Warburg Effect, that is, characteristic of cancer cells. The Warburg phenomenon is characterized by their preference for glycolysis to oxidative phosphorylation for ATP generation, irrespective of adequate oxygen supply. This is often in concomitance with enhanced glucose uptake and leads to increased lactate production and anabolic processes such as lipid synthesis and de novo nucleic acid synthesis. The molecular underpinnings of the Warburg Effect are still poorly understood. These important differences between cancer and normal cells have induced interest in glucose metabolism as a drug target. This chapter focuses on the influence p53 exerts on glucose metabolism as well as on the implications of the Warburg phenomenon in carcinogenesis and a review of the ever-increasing number of p53 regulators.",signatures:"Monde Ntwasa and Ubanako Njende",downloadPdfUrl:"/chapter/pdf-download/60555",previewPdfUrl:"/chapter/pdf-preview/60555",authors:[{id:"214732",title:"Prof.",name:"Monde",surname:"Ntwasa",slug:"monde-ntwasa",fullName:"Monde Ntwasa"},{id:"214739",title:"Mr.",name:"Phil",surname:"Ubanako",slug:"phil-ubanako",fullName:"Phil Ubanako"}],corrections:null},{id:"60895",title:"An Overview of Cancer Treatment Modalities",doi:"10.5772/intechopen.76558",slug:"an-overview-of-cancer-treatment-modalities",totalDownloads:3024,totalCrossrefCites:28,totalDimensionsCites:55,hasAltmetrics:0,abstract:"Cancer is a global issue majorly affecting developing countries. According to a survey, 63% of deaths due to cancer are reported from developing countries. There are different conventional treatment modalities that are available to treat and manage cancer. However, new cancer treatment options are being explored continuously as over 60% of all current experimental trials worldwide are focusing on tumor cure. The success of treatment depends upon the type of cancer, locality of tumor, and its stage of progression. Surgery, radiation-based surgical knives, chemotherapy, and radiotherapy are some of the traditional and most widely used treatment options. Some of the modern modalities include hormone-based therapy, anti-angiogenic modalities, stem cell therapies, and dendritic cell-based immunotherapy. This chapter discusses different traditional and novel treatment modalities to combat different types of cancer.",signatures:"Zaigham Abbas and Sakina Rehman",downloadPdfUrl:"/chapter/pdf-download/60895",previewPdfUrl:"/chapter/pdf-preview/60895",authors:[{id:"214546",title:"Dr.",name:"Zaigham",surname:"Abbas",slug:"zaigham-abbas",fullName:"Zaigham Abbas"}],corrections:null},{id:"63274",title:"Biomarker-Based Targeted Therapeutics",doi:"10.5772/intechopen.78377",slug:"biomarker-based-targeted-therapeutics",totalDownloads:1329,totalCrossrefCites:3,totalDimensionsCites:4,hasAltmetrics:0,abstract:"Cancer biomarkers are emerging as important tools for disease diagnosis, prediction and prognosis. A significant number of studies have been reported in the field of biomarker discovery due to their potential as personalized targeted therapy. With the converging gap about their utilization as specific targets, studies have focused on identifying disease-specific biomarkers in different cancer types. This chapter provides a comprehensive overview about different cancer-associated biomarkers, their prevalence in different cancer types and their use as targeted therapy. Additionally, we provide an in-sight on the therapeutic and diagnostic potential of different noncoding RNAs as cancer biomarkers.",signatures:"Shainan Hora, Amit Kumar Pandey and Sudhakar Jha",downloadPdfUrl:"/chapter/pdf-download/63274",previewPdfUrl:"/chapter/pdf-preview/63274",authors:[{id:"190426",title:"Dr.",name:"Sudhakar",surname:"Jha",slug:"sudhakar-jha",fullName:"Sudhakar Jha"}],corrections:null},{id:"62323",title:"Targeting the Ubiquitin Proteasome System in Cancer",doi:"10.5772/intechopen.76705",slug:"targeting-the-ubiquitin-proteasome-system-in-cancer",totalDownloads:2430,totalCrossrefCites:3,totalDimensionsCites:4,hasAltmetrics:0,abstract:"The ubiquitin proteasome system is involved in a myriad of biological functions including cell cycle progression, intracellular signaling and protein degradation. As such, it is not surprising to find many components of the system misregulated in cancer. The clinical success of Bortezomib for treatment of multiple myeloma proves that targeting the ubiquitin proteasome system is valid and feasible. Here, a detailed examination of the strategies used to target the ubiquitin proteasome system in cancer is discussed. The inhibitors available, its targets, the cancer type and the developmental stage it is in are discussed.",signatures:"Nishi Kumari, Kwok Kin Lee and Sudhakar Jha",downloadPdfUrl:"/chapter/pdf-download/62323",previewPdfUrl:"/chapter/pdf-preview/62323",authors:[{id:"190426",title:"Dr.",name:"Sudhakar",surname:"Jha",slug:"sudhakar-jha",fullName:"Sudhakar Jha"}],corrections:null}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"},subseries:null,tags:null},relatedBooks:[{type:"book",id:"5793",title:"Novel Implications of Exosomes in Diagnosis and Treatment of Cancer and Infectious Diseases",subtitle:null,isOpenForSubmission:!1,hash:"24e328fe01c47071f2ea44b2608e824f",slug:"novel-implications-of-exosomes-in-diagnosis-and-treatment-of-cancer-and-infectious-diseases",bookSignature:"Jin Wang",coverURL:"https://cdn.intechopen.com/books/images_new/5793.jpg",editedByType:"Edited by",editors:[{id:"188127",title:"Prof.",name:"Jin",surname:"Wang",slug:"jin-wang",fullName:"Jin Wang"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"7250",title:"Cancer Survivorship",subtitle:null,isOpenForSubmission:!1,hash:"cf8054394c93ff635e50eb4ac8cc8d3a",slug:"cancer-survivorship",bookSignature:"Dil Afroze",coverURL:"https://cdn.intechopen.com/books/images_new/7250.jpg",editedByType:"Edited by",editors:[{id:"244441",title:"Prof.",name:"Dil",surname:"Afroze",slug:"dil-afroze",fullName:"Dil Afroze"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"6813",title:"Cancer Prognosis",subtitle:null,isOpenForSubmission:!1,hash:"003e408f4cf707dd4bbf3332fe49eeb0",slug:"cancer-prognosis",bookSignature:"Guy-Joseph Lemamy",coverURL:"https://cdn.intechopen.com/books/images_new/6813.jpg",editedByType:"Edited by",editors:[{id:"182568",title:"Dr.",name:"Guy-Joseph",surname:"Lemamy",slug:"guy-joseph-lemamy",fullName:"Guy-Joseph Lemamy"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"8002",title:"Tumor Progression and Metastasis",subtitle:null,isOpenForSubmission:!1,hash:"db17b0fe0a9b6e80ff02b81a93bafa4e",slug:"tumor-progression-and-metastasis",bookSignature:"Ahmed Lasfar and Karine Cohen-Solal",coverURL:"https://cdn.intechopen.com/books/images_new/8002.jpg",editedByType:"Edited by",editors:[{id:"32546",title:"Dr.",name:"Ahmed",surname:"Lasfar",slug:"ahmed-lasfar",fullName:"Ahmed Lasfar"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"7054",title:"Current Trends in Cancer Management",subtitle:null,isOpenForSubmission:!1,hash:"0232a5ce1df00d20fe0f0189595886e4",slug:"current-trends-in-cancer-management",bookSignature:"Liliana Streba, Dan Ionut Gheonea and Michael Schenker",coverURL:"https://cdn.intechopen.com/books/images_new/7054.jpg",editedByType:"Edited by",editors:[{id:"92199",title:"Dr.",name:"Liliana",surname:"Streba",slug:"liliana-streba",fullName:"Liliana Streba"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"6143",title:"Cancer Management and Therapy",subtitle:null,isOpenForSubmission:!1,hash:"a4c510bac10f9d226b66c3559578e011",slug:"cancer-management-and-therapy",bookSignature:"Amal Hamza and Neveen Salem",coverURL:"https://cdn.intechopen.com/books/images_new/6143.jpg",editedByType:"Edited by",editors:[{id:"188326",title:"Associate Prof.",name:"Amal",surname:"Hamza",slug:"amal-hamza",fullName:"Amal Hamza"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"10477",title:"Pheochromocytoma, Paraganglioma and Neuroblastoma",subtitle:null,isOpenForSubmission:!1,hash:"ea4b534c4c57be0eaa9c5624c7e2b139",slug:"pheochromocytoma-paraganglioma-and-neuroblastoma",bookSignature:"Pasquale Cianci, Enrico Restini and Amit Agrawal",coverURL:"https://cdn.intechopen.com/books/images_new/10477.jpg",editedByType:"Edited by",editors:[{id:"196218",title:"Dr.",name:"Pasquale",surname:"Cianci",slug:"pasquale-cianci",fullName:"Pasquale Cianci"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"10437",title:"Lung Cancer",subtitle:"Modern Multidisciplinary Management",isOpenForSubmission:!1,hash:"ef76300610ffb907be22bc3e2fc6c4b3",slug:"lung-cancer-modern-multidisciplinary-management",bookSignature:"Henry S. Park",coverURL:"https://cdn.intechopen.com/books/images_new/10437.jpg",editedByType:"Edited by",editors:[{id:"96610",title:"Dr.",name:"Henry S.",surname:"Park",slug:"henry-s.-park",fullName:"Henry S. Park"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"10925",title:"Our Journey Beyond Sunset Boulevard",subtitle:"Evidence-based Analysis of Tumor-Targeted Gene- and Immuno-Therapies Shine a Critical Spotlight on “True” Long-Term Cancer-Free Survival",isOpenForSubmission:!1,hash:"dfa58123fb73c9a32fc344a3683d2126",slug:"our-journey-beyond-sunset-boulevard-evidence-based-analysis-of-tumor-targeted-gene-and-immuno-therapies-shine-a-critical-spotlight-on-true-long-term-cancer-free-survival",bookSignature:"Erlinda M. Gordon and Frederick L. 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Gordon, MD, developed DeltaRex-G (formerly Rexin-G) from bench to bedside, establishing Cyclin G1 blockade (dnG1, silver bullet) as a singular, pivotal, and strategic locus for applied/targeted cancer gene therapy. Dr. Hall served progressively as Director of Research in the departments of orthopedic, cardiothoracic, and colorectal cancer surgeries at the USC Keck School of Medicine; former President, CEO, and CSO of Epeius Biotechnologies; acting CSO of the Aveni Foundation rescue mission; current partner in Counterpoint Biomedica and Delta NextGene supportive biotechnology firms. 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He joined the staff of Cairo University (1997) and was appointed a research Professor of potential field methods in the Department of Geophysics (2014). He has undertaken affiliated post-doctoral visits to Strasbourg University, France (2018-2019), Charles University in Prague, Czech (2014-2015) and Western Michigan University, USA (2006-2007). He has authored more than 70 technical papers and served as an Editor and external reviewer for many top journals. He attended several International Geophysical Conferences in USA, Australia and France. He was a member in SEG, AGU, AAPG, EAGE and EGS. Also, he is a member of the National committee for Geodesy and Geophysics, Academy of Scientific Research and Technology, Egypt (2020-2023) and member of the Petroleum and Mineral Resources Research Council, Sector of Quality Councils, Academy of Scientific Research and Technology, Egypt (2018-2021). 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Venkateswarlu",coverURL:"https://cdn.intechopen.com/books/images_new/371.jpg",editedByType:"Edited by",editors:[{id:"58592",title:"Dr.",name:"Arun",surname:"Shanker",slug:"arun-shanker",fullName:"Arun Shanker"}],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:"314",title:"Regenerative Medicine and Tissue Engineering",subtitle:"Cells and Biomaterials",isOpenForSubmission:!1,hash:"bb67e80e480c86bb8315458012d65686",slug:"regenerative-medicine-and-tissue-engineering-cells-and-biomaterials",bookSignature:"Daniel Eberli",coverURL:"https://cdn.intechopen.com/books/images_new/314.jpg",editedByType:"Edited by",editors:[{id:"6495",title:"Dr.",name:"Daniel",surname:"Eberli",slug:"daniel-eberli",fullName:"Daniel Eberli"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"57",title:"Physics and Applications of Graphene",subtitle:"Experiments",isOpenForSubmission:!1,hash:"0e6622a71cf4f02f45bfdd5691e1189a",slug:"physics-and-applications-of-graphene-experiments",bookSignature:"Sergey Mikhailov",coverURL:"https://cdn.intechopen.com/books/images_new/57.jpg",editedByType:"Edited by",editors:[{id:"16042",title:"Dr.",name:"Sergey",surname:"Mikhailov",slug:"sergey-mikhailov",fullName:"Sergey Mikhailov"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"69579",title:"Developing and Evaluating Educational Programs",doi:"10.5772/intechopen.89574",slug:"developing-and-evaluating-educational-programs",body:'\nMost educational endeavors (e.g., producing curricula, programs, courses) follow a pretty standard set of activities for the purpose of educating students as shown in Figure 1. The chapter will rely mostly on college and university curriculum examples, but this does not exclude primary and secondary schools. Think of this as a roadmap. Like any roadmap, it is not the only way to get from Point A to Point B, but it will show the landscape and road signs from which to navigate through the process of creating new and better educational experiences for students. This chapter will briefly describe the first four phases of the process, and then focus in more detail on the evaluation phase. The emphasis on the evaluation phase is in line with current trends in education that view student learning and success as essential to academic performance.
\nSequence of activities in educational planning, designing, developing, implementing and evaluating programs and courses.
Planning is the first of five phases in creating an educational experience (e.g., an individual course, an academic program). Planning includes a set of data gathering and assessment activities aimed at helping to decide whether or not to proceed to the design, development, implementation, and evaluation phases. The output of the planning phase should be a written concept proposal that makes an academic case for proceeding to the subsequent phases. This planning document can then be submitted to the appropriate approval structures (e.g., principal, department chair, dean, academic committee).
\nA planning document needs to cover several areas, including but not limited to, a mission statement, a needs analysis, required resources, benchmark assessments, general target competencies and/or outcomes, and an evaluation plan. Before proceeding to the design phase, a few words should be said about the distinction between competencies and outcomes and the evaluation plan.
\nCompetencies refer to a general set of knowledge, skills, abilities, and other personal traits (e.g., attitudes, ethics, interests) that predict behavior in a wide variety of situations. Competencies provide the student with an integrated “mental model” of the current state and evolving standards of the field: [1, 2, 3, 4, 5, 6, 7, 8]. Examples of competencies include problem solving ability, communication skills, personal and professional ethics, and values, to name just a few.
\nOutcomes come in two varieties: program learning outcomes (PLOs) and course learning outcomes (CLOs). Learning outcomes tend to be more specific than competencies, with PLOs representing broad program objectives and CLOs representing specific ways in which a particular course meets those objectives [9]. Learning outcomes should be expressed as observable, behavioral outcomes (i.e., what the student is expected to do), and typically include an action verb and a target content area. The action verb is often taken from Bloom’s taxonomy [10, 11] that ranges from low level actions (e.g., remember, understand) to high level actions (produce, construct). For example, a PLO might be: A graduate of this program will be able to evaluate research designs and construct research projects. A CLO for a research course in the program might be: At the completion of this class, the student will be able to identify the major designs from Campbell and Stanley [12]. Another CLO for the same class might be: At the completion of this class, the student will be able to create a research project using one of the Campbell and Stanley designs.
\nAn important element in any planning document is an answer to the question: What will be used as evidence that a program or course was successful? One highly researched and successful approach addressing this question comes from the model proposed by Kirkpatrick and Kirkpatrick [13]. The model identifies four evaluation levels as shown in Table 1. These levels are (a) reaction: participant satisfaction and self-assessment of learning, (b) learning: the learners’ knowledge and skill improvement, (c) behavior: transfer of learned skills to other areas (e.g., jobs or future classes), and (d) results: impact on the institutions success and improvement. Often, educators seem satisfied with only assessing the first two of these levels (reaction and learning). The last two (behavior and results), however, may be even more essential to academic performance. The last two levels go beyond just learning, and assess what students can do and how this contributes to a more general measure of educational success.
\nSample outline of an evaluation plan with Kirkpatricks’ four levels.
The design phase involves creating a general structure for later development (see [14, 15]). Completing these steps will help guide the next phase (Development). Several actions should be taken such as (a) establish time frames for the future phases, (b) specify desired competencies or learning outcomes, (c) identify learning and performance activities that demonstrate successful achievement of the competencies/outcomes, (d) set prerequisites (e.g., students taking Algebra II must have completed Algebra I, students enrolled in a college program must have a high school diploma), (e) determine the major administrative concerns, and (f) decide what data can be collected that will reflect the four Kirkpatrick levels (see Table 1 for some suggestions).
\nFor some ideas about what design actions can be taken, see the checklist below (adapted from [14]).
Who is the primary point of contact (POC)?
To whom are the applications submitted?
How will candidates and participants be kept informed?
How will prerequisites be assessed?
Who will ensure the application materials are complete?
Who reviews and approves the applications?
When and where will training be conducted?
Where will the student records (e.g., attendance, course completion, start dates) be kept?
What budget will pay for the support personnel?
How will exams be administered?
How will exams be secured?
Who will write, proctor, and grade the exams?
Where, when, and how will skills training be conducted?
What corrective action steps will be used and who will monitor this process?
What awards and or recognitions will be issued?
Who will oversee ongoing program maintenance?
What sources of data are required to assess the Kirkpatrick four levels and how will they be obtained?
The Development phase described below explains the steps required to produce an educational practice that is ready for implementation (Phase IV). The development phase can generally be carried out in two steps.
\nThis step builds on the work completed earlier under Design (identifying learning and performance activities). This is where the actual learning and training activities are generated and matched to the learning outcomes/competencies. There are two, not mutually exclusive, options for achieving this step: (a) find relevant learning and performance activities from external sources, and (b) select or develop these activities in-house.
\nProcuring the relevant activities from an external source is far less time consuming than developing them in-house. The principal disadvantage is that the learning opportunities offered by outside sources may not be entirely suitable for the curriculum (i.e., the activities may not address the outcomes and competencies in the most direct and relevant fashion).
\nSelecting and developing the learning and performance activities in-house allows for customized experiences that can target specific knowledge, skills, and abilities. Home-grown educational experiences have the advantage of being directly relevant to the outcomes and competencies identified for the curriculum. The disadvantage of this customized approach is that it can be very specialized and may require a high degree of instructional design expertise and technical skill to develop.
\nIn this step, the learning outcomes/competencies that the students are acquiring will be assessed. The process of developing tests and measures is described in the literature on testing theory and practice (see [14, 16]). The easiest approach is to locate existing tests and measures. These existing materials may come from a variety of sources including curricula from other institutions, training classes, certification programs, continuing education units, extension classes, and so forth. Below is a partial list of candidate tests and measures that can be used to assess whether students are mastering the content and meeting the outcomes and/or competencies:
Participant satisfaction ratings
Participant self-assessment of learning
Course grades
Class quiz scores
Midterm and final exam scores
Instructor ratings of class assignments
Final project/thesis evaluation
Supervisor’s assessment ratings
Peer evaluations
Self-review of functional skills
Expert ratings of oral presentations
Simulation/game scores
Skill exercise observations
Panel review recommendations
Portfolio analyses
When an adequate set of existing assessment tools cannot be located from external sources, then customize tests and measures must be developed. When developing custom-assessment items, two important criteria must be met—reliability and validity [16]. A reliable assessment is one that is consistent. A valid assessment is one that is accurate. The first criterion (reliability) is generally established by showing that the test or measure is stable over time (e.g., repeated use of class quizzes yield consistent scores). The second criterion (validity) assures that the tests or measures accurately evaluate what they are intended to appraise. There are several techniques for ensuring the validity of tests and measures, but the most common validity check is to use Subject Matter Experts (SMEs) who closely examine the tests and measures and form a consensus that these tools in fact reflect the relevant outcomes or competencies.
\nAfter a course or program has been developed, it is ready to be implemented. There is no standardized process for implementation, but educational institutions have developed and implemented initiatives across a wide variety of disciplines and there is a large body of common practices to draw from: [14, 15, 17, 18]. In general, there are at least four steps involved in a standard implementation.
\nA pilot study is a “pre-study” conducted as a dry-run prior to launching the full effort. The study should be on a much smaller scale than the full curriculum (e.g., fewer students, less costly technology, fewer classes), but still preserve the essence of the program.
\nThe results of the pilot study should be examined and lessons learned should be noted. Specifically, at least the following elements need to be reviewed and modifications made.
\nAre these the right outcomes or competencies for this curriculum? Should more be added? Should some be deleted?
\nIs the timing of events (course duration, project times, testing schedules) optimal? Where can changes occur?
\nWere the correct prerequisites identified? Should some be added? Should some be removed?
\nWas the administration of the pilot study efficient? Where were the administrative bottlenecks and glitches? How can these be improved?
\nDid the learning and performance activities produce the intended outcomes? Should new activities be added? Should some activities be discarded? Can improvements be made to the existing set of activities?
\nDid the knowledge tests and performance measures assess the outcomes and competencies of the students as expected? What adjustments should be made?
\nAre the data collected easily obtained and in a usable form? Can clear conclusions be drawn from these data?
\nTo help ensure success, a marketing plan should be devised to advertise the program and recruit students. The following items should be considered: (a) identify the target audience, (b) align the marketing objectives with the curriculum objectives, (c) create a communication plan, (d) publish a schedule, and (e) use specific institutional marketing techniques (e.g., fact sheets, web and electronic media, newsletter, brochures, communication networks, open house events, personal visits to potential recruiting venues).
\nIn this step, the program gets implemented in accordance with the pilot study modifications.
\nAfter the course or program has been implemented, it must be evaluated for effectiveness. This evaluation should be driven by some formal model such as the Kirkpatrick and Kirkpatrick model [13] shown in Table 1. If the Kirkpatrick’s model is adopted, then data are required that assess each of the four levels. If the evaluation is for a single course, then the tests and measures will be mainly, but not exclusively, relevant to level 2 (learning). If the evaluation is for an entire program, then all the levels should be assessed (as shown in Figure 3 to be discussed below). I’ll begin with the evaluation of a single course, and show one possible approach.
\nOnce the knowledge tests and performance measures have been administered to students in a class, each person should have a set of relatively objective scores. These scores, when combined, should show how successful the student was with regard to the class outcomes. The approach illustrated here is based on “performance indexing” developed by Tatum and Nebeker [19]. Performance indexing is a system for combining and weighting a set of scores and generating an overall index. Performance indexing has been employed successfully in fields outside of education (e.g., real estate, environmental quality management, organizational improvement), but can be used just as effectively in an educational setting. The weighting feature is especially useful because it takes into account how valuable each test or measure is in the overall assessment. If, for example, in a biology class, mid-term and final exam performance is more important than homework, this difference will be reflected in the final index. Often, the degree of importance is reflected by the number of points that can be earned by each assignment. Performance indexing offers a more sophisticated system for balancing performance and getting at the essence of student learning. An example of performance indexing used in a hypothetical class is shown in Figure 2.
\nA performance index table used to assign a final course grade to a single student in a hypothetical class.
There are several steps to developing and using performance indexing (for a more complete discussion of the topic see Tatum and Nebeker [19]). The most essential features are (a) each test or measure is given a weight according to its importance in the assessment, and (b) an overall index score is generated from the weighted values (e.g., 370–400 is outstanding performance). Table 2 is a step-by-step guide for building and using the performance index table in Figure 2.
\nSteps for building and using a performance index table.
A program (e.g. clinical psychology, biology, history) is designed so that students graduate having met certain competencies or PLOs. Whether the program uses competencies or PLOs is a matter of preference, but regardless of this choice, the CLOs must be designed to meet one or more of these competencies or PLOs. When students successfully complete all of courses in the program, they will have satisfied the expected objectives of the program and will leave with specific knowledge, skills, abilities and other desired characteristics (e.g., attitudes, personal ethics, interests).
\n\nFigure 3 is an example of how performance indexing can be used to evaluate an entire program (as opposed to a specific course within that program as shown above in Figure 2). Developing an index table for a program involves basically the same steps outline in Table 2, with a few modifications. The evaluation measures (shown in the diagonal spaces) are based on the Kirkpatrick and Kirkpatrick [13] levels (see Table 1), which are shown at the top of the Figure 3. The specific evaluation measures will vary from program to program, but each measure should fall into one of the levels. For example, level 1 (reaction) is supposed to indicate the students’ satisfaction rating of the program and an assessment of how much they think they have learned. These ratings can be obtained from each class or as part of an exit survey at the end of the program. Level 2 (learning) is intended to reveal, on a more objective basis, how much the students learned (in this case based on grades, ratings of acquired skills, and test scores average across classes). Level 2 is closely tied to the competencies or PLOs of the program. Level 3 (behavior) is an indication of the degree to which the program changed the student’s behavior and the extent to which the student can transfer this behavior to other settings (e.g., did they learn valuable job skills, did the acquire knowledge and skills in prerequisite classes that they can apply to future classes?). Although level 3 is normally associated with assessing an entire program, it is still possible to include behavioral measures at the course level. For example, Figure 2 shows an “internship supervisor rating” as an essential measure of the student’s ability to apply what was learned. Finally, level 4 (results) is supposed to show that the program had a positive impact on the current success and future improvement of the institution (e.g., local school, school district, college). Evidence for positive results can be demonstrated by a variety of data such as graduation rates, employment success, advancement to higher levels of education, or ratings by external agencies. Level 4 measures are not common among educational institutions when evaluating individual programs (although these data are routinely collected at higher levels), but they should be. To capture the essence of academic performance, we must assess the degree to which our programs contribute to the general success and welfare of the broader academic community.
\nA performance index table used to evaluate a hypothetical academic program.
Once the final index is computed, the overall success of the program can be evaluated. In the hypothetical program depicted in Figure 3, the index score of 300 indicates that the program is above average. A close examination of Figure 3 will also reveal where the program is performing well (students rate their learning and job skills as exceptional, the program gets an exceptionally high rating by external agencies) and where it requires improvement (test scores are low, there is a low percentage of students finding jobs or advancing to other programs).
\nThe phases and steps advocated here are obviously a mechanistic (non-theoretical) approach. It resembles Tyler’s [20] thinking about curricular design more than contemporary thought (e.g., [21, 22]). There is nothing wrong with a more mechanical approach. In fact, the phases and steps proposed in this article are not incompatible with modern views of education such as the sharing of common goals [23], scaffolding [24], or the spiral curriculum [25]. At some point, however, we need to find and follow a path towards building an educational program, and this roadmap shows us the way without too many detours.
\nAcademic performance has been the focus of much research and interest during the past few years. Initiatives such as No Child Left Behind [26] and Race to the Top [27] have generated much debate and concern regarding the components of academic performance [28] and the optimal methods for assessing learning and success [29]. This chapter proposes a method for developing and evaluating courses and programs that gets at the heart of academic performance in five phases (i.e., planning, design, development, implementation, and evaluation). The first four phases are a prelude to what the author considers the true essence of academic performance; namely, the identification and measurement of performance indicators. This chapter presents an evaluation model (based on [13, 19]) that guides the user down a well-traveled road that leads, in the end, to a quantitative understanding of student course performance and program success. In the inimitable words of Peter Drucker:
The author would like to acknowledge the valued assistance of Dr. Don T. Sine for his support and critique of parts of this research and manuscript.
\nTrophic relationships between organisms are the mechanisms responsible for most of energy and nutrient transfers; they allow the functioning of the ecosystem. These relationships, known as food webs, caught the attention of naturalists before the concepts of evolution and ecology were about to be determined.
Initially, the diet of a species and its skills to obtain it were recognized as the leading factors for the prevalence of the fittest. Additionally, it is one of the main forces leading to evolution of that species in the long term [1]. Furthermore, competition for food became one of the favorite hypotheses to explain species exclusion; it states that when two species seemed to feed on the same resources, the best suited ultimately leads its competitor to extinction in the long term [2]. This idea has been around for many years and has not been completely discarded or proved [1].
Examining phototrophs, also known as primary producers, is the dominant starting point to analyze food webs. They use the incoming sun’s energy and inorganic nutrients to generate their biomass. This is the most important mechanism, as it initiates the cycling of nutrients and energy flux in aquatic food webs. There is primary productivity involving chemolithotrophs dominating in places devoid of sun’s light [3]. These places were mostly known to be, until recent times, around underwater volcanoes more than 1000 meters deep [3, 4].
Primary production is at the base of all consumers concurring in the environment. However, macroscopic food webs tend to be very short, with few levels of consumers because these organisms dissipate matter and energy efficiently [5]. All metazoans invest their energy looking for food, ingesting it, digesting, repairing themselves, mating, and reproducing. These activities make multicellular organisms to get around 10% of biomass fixation efficiency. Thus, 1,000 kgs of the primary producer will be needed to produce 100 kgs of herbivorous animals, only 10 kgs of small carnivores, 1 kgs of medium-sized carnivores, and only 0.1 kgs of top carnivores, following a pattern known as pyramid of productivity [5]. Adding a predatory species at any level would destabilize the food web, as this will consume higher amounts of biomass [6]. Energy dissipation is even larger, meaning that the entropy produced during the functioning of the food web is very high. However, only 1% of the incoming sunlight is used for primary production, stressing the importance of the environmental factors limiting biomass productivity to sustain food webs.
Primary productivity varies along seasons. When it reaches its peak, productivity is controlled by the top predator’s consumption (top-down), and when it reaches its lowest level, productivity is controlled by phototrophs (bottom-up). There are places that are permanently bottom-up controlled such as the deep ocean communities depending on the “organic matter rain” from dead organisms living in the photic zone in places near the equator are almost always top-down controlled, where productivity may be at its peak for most of the year. All other places experience top-down/bottom-up controls alternatively, depending on the productivity seasons.
Unicellular algae lead primary productivity in marine environments, sustaining the great diversity of organisms, especially in places receiving nutrient inputs from lands. Heterotrophic unicellular organisms forage on algae and both phototrophs (phytoplankton) and heterotrophs (zooplankton) conform to the plankton. However, unicellular organisms span in sizes less than 1 μm to hundreds of micrometers, and the species’ diversity of plankton, including microbial eukaryotes and bacteria, ranges in the order of thousands. Species of microorganisms are much more numerous than the metazoans. With such a great diversity of microorganisms, it become apparent that the microbial food webs may function differently from the macroscopic food webs.
It was believed that food webs would get destabilized if the number of species increases at any level above the primary producers. However, microbial food web seemed to get more stability with the increasing number of species, contradicting what was observed in macroscopic food webs [7]. Thus, the higher number of species of bacterial and microbial eukaryotes in aquatic food webs seemed to contradict that assumption; this phenomenon was named as “The paradox of microbial loop.” It was paradoxical that productivity and efficiency of nutrients and energy transformation is increased by adding more species, promoting the stabilization of the food web [8].
It’s been a long road since the recognition of the “paradox of the microbial loop” in the aquatic food webs. Nowadays, it is referred only as the “microbial loop,” after being integrated into the food web conceptualization in both terrestrial and aquatic environments [7].
The complexity of microbial food webs needs to be approached from the analysis of different functioning capacities and nutritional needs of the participating species. It has been normal to assign very general feeding habits to protists and metazoans, like bacterivores for example. This nomenclature implicates that a single species of protist can feed on any one or indistinctly on all the thousands of bacteria species. However, observation of feeding habits has revealed that protists and metazoans prefer feeding on specific kind of bacteria while avoiding other species. Pigmented bacteria [9], for example, has fewer predators than non-pigmented ones. On the other hand, there are several species of protists, mostly amoebae, small flagellates and
One explanation for pigmented bacteria to have fewer predators relied on the toxicity or poisonous effect of those pigments for many protists, pointing out the importance of the biochemical warfare that bacteria must synthetize to defend themselves. However, chemicals used for evading enemies attract other ones looking for those same compounds, putting bacteria in a situation where there is no way out for bacterial preys. Indeed, there is no way out of being preyed upon, as every living being has predators, or at least other species which may feed on them or use them as a resource.
Is there a single factor determining the feeding preferences? The short answer is “No.” Remember that “bacterivorous” or “algivorous” are labels used to recognize the kind of food that protists and metazoans may prefer to feed on, and it involves many species. From the beginning, this was a non-exclusive way to label the category of food that may be used to group the highest quantity of species to simplify and conceptualize the food webs. Furthermore, during the first half of the XX century [11], there were many very interesting studies trying to determine the “diets” of several species of protists [11, 12], with the aim of designing a chemically defined culture media, as is the case of several recipes for culturing
Designing a culture media for protists or bacteria was a major task, as numerous factors about their nutritional needs were unknown (and remain unknown). These attempts to cultivate bacteria and protists lead to one important conclusion: different species cannot synthesize one or several molecules needed for their metabolism and have to take those molecules, as such, from their ingested food [12] or from other microorganisms that live within the biofilm (such as the case of NAD+ **, which the bacteria has to consume from other species of bacteria for both of them to grow). Microbial biologists named this phenomenon as “auxotroph” [13]. In this way, the molecule(s) a bacterial species is auxotroph for must be added to the culture media, to keep a culture of such species [14]. The kind of molecules, their diversity, and their macro- and micronutrient composition form a universe comparable to the one containing the species’ diversity on the planet.
Ecological relevant functions have been recognized in prokaryotes and microbial eukaryotes. Bacteria have been cataloged as nitrogen fixing, denitrifiers, metanogens, methanotrophs, phosphorous mobilizers, metal mobilizers, phototrophs, and chemolithotrophs as the main recognized functions in the ecosystem. On the protists’ side, several trophic groups have been recognized as phototrophs and phagotrophs. The first group is strictly divided between the phototrophs and mixotrophic ones, while the second one may be divided in bacterivorous (including cyanobacteria), frugivorous (feeding on hypha and or yeasts cells), algivorous, protist consumers (raptorial protists), and metazoan predators. Parasitic bacteria, pathogenic bacteria, and microbial eukaryotes have been largely studied from the medical point of view. However, recently, they have been studied from the ecological perspective (their impact on the predator–prey relationships, the “health” of species populations protected for conservation, and their effect on the nutrients distribution along food webs [13].
Phagotrophic protists may ingest very different kinds of particles and present the capacity to eject the ones they cannot digest, or even reject particles previously ingested [15]. Even if the water current would transport a good mixture of different bacterial species, phagotrophs may choose which particles ingest and eject the debris from their digestion together with the non-digestible microorganisms. This means that protists may show preference for the kind of food they most likely can digest (recognizing their preys by their quorum sensing signals), and, like bigger organisms, they may need a variety of food sources to get the nutrients they need [15].
A close examination of the different trophic groups allows to re-mark the unicellular phototrophs as the most productive in terms of biomass production since there is no synthesis of support or conductive structures, and, because of that, they are the base of the aquatic food webs.
The phagotrophic protists have been recognized for being the main consumers along microbial trophic networks in aquatic systems conforming a major proportion of the microbial biomass in these systems [16, 17]. These predators are also responsible for much of the recycling flow of nitrogen and phosphorus in the aquatic systems [18].
Particularly the ciliates are key elements of aquatic food webs they have several functions, they can be primary producers, predators, they serve as food for metazoans including free-living stages of metazoan parasites; there are many aquatic habitats without macro-organisms, but none without bacteria and at least few protist species [19].
One of the most interesting groups of protists are the mixotrophic ones. Some of them may correspond to the old morphological groups of ciliates, flagellates, and ameboebas. Mixotrophy is defined as the ability to combine phagotrophy and phototrophy in a single cell [20]. This group can be divided into constitutive mixotrophs, meaning they have the innate ability to photosynthesize, and the facultative or non-constitutive mixotrophs. These organisms may sequester the plastids after consuming their phototrophic preys or by harboring photosynthetic endosymbionts [20, 21]. Around 23% of planktonic ciliates species (marine and freshwater combined) perform acquired phototrophy, and this ability is present in at least 8 main ciliated taxa: Heterotrichea, Hypotrichia, Oligotrichida, Stichotrichida, Litostomatea, Prostomatea, Peniculia, and Peritrichia. Phototrophy is usually acquired from algae endosymbionts in 7 of these 8 ciliated taxa. Contrastingly, Oligotrichida usually obtains this ability by plastid sequestering [22].
The structures of the mixotrophic ciliates community varies through seasons, depending on the changing water trophic condition. Mixotrophic ciliates dominate in spring and summer, reaching from 58–100% of the ciliates in oligotrophic waters [23, 24, 25], but represent only 5% of the total community of ciliates in winter, probably due to the lower water temperatures and nutrients. These conditions restrict the growth of algae, negatively affecting the population of mixotrophic ciliates if their preferred species of algae is missing [24].
The mixotrophic ciliates are mainly from the genera
Ward and Follows [33] performed a global simulation of the ocean-surface food web, revealing that mixotrophy enhances the transfer of biomass to larger organisms at higher trophic levels, which in turn increases the efficiency of oceanic carbon storage through the production of larger and faster sinking conglomerates of organic molecules. It follows that mixotrophic protists play a key role in modulating the primary production that underlies the food web in aquatic systems [21, 22, 32]. However, their importance has not been fully appreciated because traditional field and laboratory studies focus on strict classifications as phototrophs or phagotrophs [32] because incorporating this flexibility to acquire food is difficult to modelize. Mixotrophy is known to be common in all aquatic systems but its contribution to net community production is difficult to quantify, and the integration of their impact on the global biogeochemical cycles remains to be incorporated.
Ciliates and flagellates are the most dominant bacterivores among the phagotrophic protists in most aquatic systems [16, 34], consuming between 25–100% of the daily production of marine phytoplankton together with large quantities of bacterial biomass [18]. Bacterivores and algivorous protists are the core consumers of microbial biomass in aquatic food webs [16, 17] regulating these groups in two apparently contradictory ways: by feeding on the abundant food source, they keep in check their further expansion, that in turn gives other less preyed species the opportunity to become more numerous, and at the same time, the release of cellular wastes (from protists) enhance the reproduction of the species being predated. The combined effect of these two processes enhances the nutrient cycling and fuels biomass productivity. By performing this activity, ciliates and flagellates increases their own biomass, attracting metazoan predators and functioning as linkage of lower and upper trophic levels in aquatic food webs [16, 35, 36].
The size of the ciliate determines the sizes of preys they can feed on. Thus, bacterivorous ciliates ingest a different particle size range; the preferred size spectrum for each species is a function to cytostome size and morphology. For example, small ciliates usually bacteria eat <3 μm [18, 37, 38]. Ciliates that feed on the smallest particles (<1 μm) require relatively high densities of these bacteria as a minimum to keep their population growth [30]. Several groups of ciliates actively feed on specific bacteria species for a period ranging between 44% and 100% of the time, because bacterial densities will have variations as responses to predation intensity along time [36].
Bacterivorous ciliates are present in all aquatic environments, from oligotrophic to eutrophic, in both freshwater and oceans. The diversity of bacterivorous ciliates and their contribution to the flow of energy in trophic networks depend on the dynamics of the systems in which they are living. Therefore, food resources are probably the main regulators of ciliated communities (diversity, abundance, and biomass) [30]. For example, bacterivorous ciliates contribute very little for the direct transfer of bacterial production to the trophic networks of metazoans in oligotrophic environments. Ciliates consume less than 11% of bacterial productivity in these waters [39, 40, 41]. Perhaps the heterotrophic bacteria that are very small in these lagoons (0.035 to 0.4 μm) are grazed by bacterivorous ciliates at a very low rate [41], or the number of bacteria is not enough to support larger ciliate communities feeding on smaller bacteria (<1 μm), as they require high densities of bacteria to maintain their populations [30]. Then, productivity of oligotrophic systems function most of the time as bottom-up (availability of substrate and nutrients) controlled [42]. This functioning will remain until seasonal pulses of nutrients (or human subsidies) arrive, busting primary productivity and changing the system into top-down control, and it will keep functioning the same way until the pulse of nutrients (or subsidy) is completely metabolized, returning the system to the bottom-up dynamic.
Contrastingly, densities of heterotrophic bacteria in eutrophic environments are sufficiently higher to also keep a higher diversity of active bacterivores [43], fueling ciliates biomass productivity and allowing the intervention of metazoan predation. Top-down control (predation) seems to be in function all the time for regulating the abundance of heterotrophic bacteria in eutrophic systems [42]. Normally, communities of bacterivorous ciliates of small sizes (~ 30 μm) are found as dominant in eutrophic environments [30, 38]. The most abundant ciliates in these environments are small oligotrichs (
Sessile ciliates such as
Trophic groups | Examples | References |
---|---|---|
Bacterivores | Colpodida ( | [34, 46, 48, 114, 116] |
Feeding on Phototrophs | Choreotrichia ( | [54, 56, 57, 58, 60] |
Predators of predators | Heterotrichea ( | [62, 64, 66, 68, 72, 108] |
Omnivorous | Choreotrichia ( | [49, 66, 69, 73, 79] |
Mixotrophos | Litostomatea ( | [23, 25, 26, 28, 29, 31] |
Trophic groups free-living ciliates in aquatic environments.
There is a difficulty in assessing a proper name for the kind of food protists feed on when they become predators of phototrophs, as this group consists of both eukaryotic and procaryotic members, and neither of these primary producers may be considered as “plants” or “herbs”. Feeding on them cannot be considered as herbivory. On the procaryotic part, cyanobacteria are a phylum comprising many species that, besides being phototrophs, can also produce toxic molecules, compromising the fresh water supplies for human consumption when growing unchecked in oligotrophic waters [50, 51]. From the eukaryotic part, there is an extra complication when trying to separate the permanent phototrophs from the mixotrophs.
Moving the sizes up, ciliates are one of the most important groups feeding on phytoplankton in marine and freshwater environments [18, 41, 52]. They may consume up to 74% of the daily phytoplankton production [53], becoming the key controllers of phytoplankton biomass [54]. On the other hand, ciliates mobilize the highest proportion of organic carbon and nutrients in oligotrophic waters dominated by cyanobacteria, playing the fundamental role of linking the productivity of microbial food web with the metazoans [41, 53]. It has also been noticed that the flux of carbon up to metazoans is not interrupted when the density of bacterivores ciliates falls, but it is compensated by predation on ciliates feeding on phototrophs [41]. Some of the ciliates that feed on phototrophs are in Table 1.
Ciliates feeding on phototrophs represent between 30–65% of the total biomass of all functional groups of ciliates thriving in eutrophic lakes [55]. However, this dominance is not permanent. Ciliates feeding on phototrophs become very numerous on the blooming season [56], and even dominate the entire ciliate community for short periods between seasons [57].
Tintinnids tend to feed on small-cell-sized phytoplankton (2–20 μm) [58]. They are voracious phytoplankton feeders that may consume over half the quantity of these kind of phototrophs in marine waters [54] and over 69% of these primary producers in lakes [59]. Species like
Selective feeding has been observed in several species of ciliates. However, feeding on a wider spectrum of sizes and kind of phototrophs (non-selective feeding) allows them to take advantage of the productivity in hypereutrophic environments rich in small particulates [49]. The genus
There are several species of ciliates and flagellates that feed on bacterivorous protists and on protists feeding on phototrophs. These are predators of predators. These predator species may feed temporarily on bacteria but cannot survive by just this consumption; they are attracted to them as they offer clues to discover their preferred preys: ciliates, flagellates, or amoebae feeding on bacteria.
Most of predator ciliates feed on preys around 10 times smaller than them [62, 63], although raptorial feeders may consume bigger preys, comparable to their own size or even bigger [64]. This capacity is due to their very flexible cytostome as is the case in protostomatids genera
Predatory ciliates are present in small numbers along seasons in oligotrophic waters, showing surges in population numbers, in synchrony with the increase of primary productivity during the spring, reaching up to 55% of the total ciliates’ abundances in temperate waters [64, 66]. However, they only reach between 24.6% to 28.7% in freezing oligotrophic waters of the Arctic and Antarctic [67].
On the other hand, predatory ciliates become important top-down controllers of microbial food web productivity in eutrophic and hypertrophic waters [68]. Eutrophic waters have the conditions to sustain high productivity rates of phototrophs and heterotrophic bacteria, sustaining, in turn, large populations of their grazers, promoting the increase of predatory ciliate population [69]. Biomass of raptor ciliates may reach almost an order of magnitude higher in eutrophic compared to the one obtained in meso and oligotrophic lakes, suggesting that they are effectively controlling the primary productivity [70]. This assumption is supported by the covariance of predatory ciliates and their preferred food. For example, the increasing population of predatory ciliates bigger than 100 μm is related to a simultaneous shrinkage of abundance of smaller ciliates (<20–40 μm), mostly phototrophs and bacterivorous [71]. Big and voracious ciliate raptors like
Several species of oligotrichs feed on bacterivorous flagellates, showing an efficiency of 45% biomass transformation, also fueling the bacterial productivity by releasing essential nutrients for heterotrophic bacteria to keep their population growth [65]. Some predatory ciliates are shown in Table 1.
Omnivorous protists are an important group to look for when assessing the stability of a food web because their very presence means productivity is enough to non-specialists, to feed on a variety of resources. Omnivores strengthen the resilience of planktonic communities by regulating the trophic dynamics [73]. Omnivorous ciliates may have a preferred prey but can easily move to other kinds of prey, which may be more abundant or easier to catch [74]. This variety of resources for true omnivorous ranges from bacteria, algae, other ciliates of different sizes to fungi [73]. This versatility gives them an advantage to withstand a resource limitation by having alternative prey [70]. Additionally, omnivorous ciliates increase the stability of planktonic communities by feeding on species that may pass undetected from their specialized predators, by having densities small enough to get an advantage of the elimination of their competitors and increase their numbers. In this situation, omnivores would prevent them to reach high densities too fast, giving time for their specialized predators to increase to population levels that may effectively control the newly abundant prey.
Omnivorous ciliates are present in any kind of environment allowing the stability of protist communities. They are elements of marine and freshwater ecosystems, both oligotrophic [66, 75] and eutrophic [69, 76], as well as in polar waters [67].
As with the other trophic groups, omnivores show seasonal bursts of abundances in the communities they are part of, especially in oligotrophic waters where they are scarce most of the time, except for occasional bursts [77, 78]. Omnivorous ciliates are commonly found in lakes throughout the year, normally with low species richness, representing between 2–12% of the ciliates species [67, 79]. Their low contribution to the number of individuals makes them reach a peak of 35% during productivity bursts [66, 79]. However, this proportion may steadily increase in the proportion the environment is turning into the eutrophic condition, increasing the species richness, although the densities of omnivorous ciliates may momentarily diminish with the eutrophication [69] as result of the species increase (more species and lower number of individuals by species). Once the eutrophication reaches a steady state, the biomass of the omnivorous ciliates will reach high values and even dominate among ciliates [76].
The numbers of small omnivorous ciliates usually dominate in meso oligotrophic environments, feeding on dominant bacteria (<2 μm) and algae (2–20 μm) [49]. Food concentration is a very important factor, strongly affecting an easily detectable feeding behavior of omnivorous ciliates [73]. Several of the most common omnivorous ciliates are shown in Table 1.
Functionality alone has its own complexity in food webs because, for example, mixotrophs would be functioning as phototrophs or as heterotrophs along different hours during the same day (How long do they function as phototrophs? How long do they the function as heterotrophs?). An extra dimension in this world comes from the different sizes of preys corresponding to the predators’ sizes and the number of cells each individual predator must get to produce another individual [80]. This is one of the reasons why plankton has been divided in microplankton, nanoplankton, and picoplankton. Each category corresponds to the range sizes of microorganisms. The smaller ones like picoplankton and nanoplankton, performing primary productivity (chemolithotrophs or phototrophic [3], can sustain their corresponding predator’s size and be up to ten times bigger, namely nanoflagellates and microflagellates. These are the two groups of protists related to their size and morphology rather than their taxonomic affiliation [81], since very few information is known about them apart from 18S SSU rDNA sequences; they have been recognized performing predatory activity on phototrophs of the smaller sizes.
One alternative to conceptually reduce the complexity of microbial food webs is analyzing them as nested compartments. This means that the transfers of matter and energy takes place inside each compartment corresponding with one size class of producers and its predators because these organisms function in the same time frame. Then, several of these compartments may get integrated in a bigger one by predation of the next size class. Time frame for this bigger class is also bigger than the previous one, as the sizes of the organisms are also bigger and so on. Every compartment of bigger sizes function as concentrator of biomass and disperser of energy. However, the wastes generated in each compartment releases the nutrients once fixed in the biomass fueling the nutrient cycle in compartments of all sizes. Up to here, it looks like the aquatic food web is functioning as a continuum along and through the water column and surface. However, there is a chance of recognizing boundaries to help a better understanding the food webs dynamic.
When hearing the word “boundary”, immediately, the existence of physical barriers delimiting something in space comes to mind. Because of that, it is hard to imagine an aquatic food web being physically limited because our experience has shown us the big animals feeding on all planktonic organisms at once, which could be in thousands or even millions. However, it just represents a small appetizer for a whale.
A careful examination reveals that very small organisms live faster than ones at the immediate upper-sized scale and intuition tells us that time may be experienced in different ways, depending on the size of organisms involved. The size ranges occupied by ciliates in the microbial food web spans from less than 10 μm to more than 4500 μm [82]. Comparatively, their pool of size ranges would be like the pool of sizes from small fishes to whales. Why are these sizes important? Because it can be argued that the velocity of nutrient exchange is faster in the smaller organisms and the nutrients may be “sequestered” for long periods by the bigger and long-lasting animals. In this way, a complication of time arises when trying to diagram the nutrient cycle in the microbial food web. Time becomes another varying feature rather than a constant in food web dynamics. In this way, time may draw the boundaries between compartments and, at the same time, could be avoiding contradicting the nested compartment proposal in the physical limitless aquatic system.
It is easier to recognize physical boundaries in terrestrial ecosystems as the environment changes at slower velocities than the very dynamic aquatic environment. Soil is a heterogenic environment, the opposite to the aquatic ones. It is an environment that cannot be seen through and be dived in. Soil matrix is composed of a very complex mixture of mineral particles, organic matter and living organisms. This mixture is organized in aggregates that may facilitate or resist water and air passing through it but, most importantly, these aggregates proportionate spaces where all living beings can move through soil.
At a microscale, soil aggregates divide the open spaces in two types, the fast water passing by (the space between aggregates) and the slow motion of water in the space inside the aggregate, and consequently of slow-moving air too, as air and water move through the same spaces). These are the soil’s physical boundaries, and this is the environment where roots move and look for hotspots of nutrients, as well as places where microbial symbionts may be found (normally inside soil aggregates). Water reaching soil aggregates dissolved salts and polar molecules that may contain nutrients that will be taken by roots, mycorrhiza, or bacteria. This is a complementary start of plants primary productivity, because plants have to take water from soil together with other nutrients to produce a wide range of molecules, from non-protein forming amino acids to scents and pheromones, as result of what is known as the “secondary metabolism.” Plant primary productivity comprises both photosynthesis-respiration (primary metabolism) and secondary metabolism, irrespective of being vascular or nonvascular.
Soil productivity is dependent on the nutrient exchange velocity rather than the gross amount of bioavailable nutrients. Nutrients used and released very fast means energy is being captured, transformed, and degraded very fast, implying the activities of all participating organisms are taking place so fast that production of biomass at all levels is gaining momentum and its control may come only from consumption (top-down) no matter that nutrients exist in limited quantity. This feature also explains why the smaller organisms can sustain productivity of the biggest ones. In other words, aerial part of plants are very important for primary productivity because it is the place were light, inorganic carbon, and water are used to produce organic molecules that are at the base of primary productivity (Sun’s energy fixation in organic molecules).
Without diminishing photosynthesis’ importance, most of terrestrial plants gather a “productivity teamwork” inside and around their roots, involving mycorrhizal fungi and mutualistic bacteria, a functional place known as the rhizosphere. Almost 80% of the known terrestrial plants need the association with a mycorrhiza, to appropriately complete their life cycle, but all plants need mutualistic bacteria to grow. Microbial partners are indeed an important part of primary productivity, as they actively participate in the acquisition, modification, and metabolization of many organic molecules containing the elements we call “Nutrients.” For example, it has largely been demonstrated that mycorrhiza translocate phosphorus to plants. At present, very few people challenge this. However, what form of phosphorus is translocated from mycorrhiza to plant? Surely, it is not the phosphorous as molecule, but organic molecule where P is forming part of the structure. Plants can take up P from inorganic molecules in general or from phosphoric acid. Why do they need mycorrhiza to supply P? It is still an open question, but the degree of specificity of the plant-mycorrhiza association allows to conjecture that plant and mycorrhiza share metabolites containing nutrients (not just P) for metabolic complementation, and the same could be true for mutualistic bacteria. This would explain why one species of mycorrhizal fungi is mutualistic to several plant species but functions as pathogenic or parasite to other ones.
Contrary to what happens in waters, soil fungi and bacteria are scattered through soil and physically constrained to available surfaces. If they keep growing unchecked, bacteria may become effective nutrient competitors to plants, as nutrients forming bacterial biomass are non-available to plants. Mycorrhiza may move farther away from the root than bacteria and can establish a mutualistic relationship with other roots (whether they are from the same plant or from a different species, it does not matter) to avoid becoming competitors. Absence of bacterivores is a needed condition for bacteria to become a plant competitor in the rhizosphere [83, 84]. Bacterivores ciliates, flagellates, and amoebae release nutrients trapped in bacterial biomass, stimulating both plant and bacterial growth. In the first case, nutrient release allows roots to take them in and bacteria microcolonies may grow again in the root surfaces, already cleaned out, and obtain nutrients from predators’ wastes [84].
Soil’s physical constrains allow growth of bacteria and fungi in differentiated places. Sometimes bacteria also grow on the surface of hypha, helping fungi to mimic bacteria and somehow escape from fungal predators. It has been possible to observe protists feeding predominantly on fungi and avoiding bacteria as much as possible (
This differentiation of soil’s physical spaces makes it easier to visualize the small productivity compartments around roots, absorbing hairs inside small soil aggregates, bigger compartments covering aggregates on the tip of the root and getting in contact through fungal hypha.
Motility of bigger protists are limited to litter and upper soil layer by the available spaces, restricting their abundance in the underneath layers. Testate amoeba, ciliates, and flagellates, around 100 μm, dominate in these 2 layers and actively participate nutrient recycling from litter, while smaller size ciliates like
Primary productivity in soil is restricted to the upper layers where cyanobacteria and eukaryote algae may survive and even form thin layers known as microbial soil crusts. Both phototrophic bacteria and algae may form stable mutualistic symbiosis with other organisms, like fungi, to develop thicker structures composing soil crusts showing lichens and mosses. Beneath and into soil crusts, ciliates, flagellates, and amoebae are among the most important microbial predators, active mainly during the time of water availability [85, 86]. However, the main photosynthetic carbon input is released by roots into soil layers [87]. Roots secrete amino acids and other complex organic molecules to attract symbiotic bacteria and mycorrhiza conforming the trio of soil productivity sustaining microbial food webs deep into soil [88, 89]. Consequently, protists’ species diversity may be higher around roots and the dominance of ciliates may be restricted to the sizes of soil pores [86, 90, 91, 92]. Soil protists were recognized as purely bacterivorous because fungi feeding protists may transitorily feed also on bacteria. However more detailed studies have recognized species of soil protists feeding only on bacteria or fungi [93, 94, 95]. Among the main bacterivorous ciliates are Colpodida (
Fungi and bacteria normally use different kind of organic molecules, bacteria normally metabolize low molecular weight organic molecules while fungi normally metabolize complex organic polymers of high molecular wight [97]. This metabolic difference allows to conceptualize two pathways for nutrient cycling: the bacterial and the fungal paths. However, this concept is being challenged because of the abundance of protists feeding on both kind of microorganisms [98, 99]. All the early recognized fungi feeding ciliates and amoebae in soil ranges from 50 microns to above 150 μm [100]. However, there are also smaller ciliates and flagellates feeding on both spores and hypha [100]. The main groups of specialized fungal feeder ciliates are grouped in the family Grossglockneriidae [93]. This family of ciliates may account for mora than 2% of the protists sequences in the forest litter and grassland while may drop below 0.3% in peatland soil, probably due to the reduction of soil pore sizes [100]. Although, counting techniques based in MPN calculated around 200 cells/gram soil DW in previous studies [101]. Protists have a very limited capacity to disperse throughout the soil system by themselves. However, oligochaeta disperse them as cysts farther than a few centimeters, in the range of several meters both horizontally as well as vertically into the soil system.
Soil functioning is much more variable than the aquatic systems, as it is regularly subjected to dryness and several flooding events per year. For microbial ecologists, soil is a natural stressed environment, having enormous variations of water availability through seasons, especially in arid and semiarid environments. However, there is a comparable situation, although at lesser degree, in the tropical dry forests, temperate, and tundra regions. Even at the equator, the rainy forests may show an excess of soil in water, stressing microbial food webs.
Microbial communities have been evolved by modifications and adaptations in responses to natural stresses that finally allow them to get along with environmental change. The problem we are facing now resides in the velocity of environmental changes imprinted by human activities. The most important, but hardly the only one, resides in the use of fossil fuels because of the acceleration of climate change. The CO2 released as byproduct of combustion is just one of the causes of climate modification in the short term (in historical and geological times). Internal combustion engines also produce other greenhouse gases such as NxO or NO2, having a bigger capacity of keep heat, and this is a big problem generated only for the atmosphere. Hydrocarbons pose a permanent threat of contamination to aquatic and soil systems near the extraction zones, the transporting infrastructure to refineries, infrastructure for later transportation as fuel to expending places, and by illegal activities damaging oil ducts.
Soil microbiota react in different ways along the gradient of contamination when hydrocarbons reach soils. The plume of contamination normally eradicates the phototrophs and exert a strong selective pressure on bacteria and fungi, by killing or inhibiting the growth of sensitive species while enhancing the growth of resistant ones. These effects can be modified by the toxicity of the different compounds rupturing and/or changing the connections of the trophic networks [102, 103].
The effect of hydrocarbon contamination and others contaminants (pesticides, heavy metals) on communities will depend on the intensity, duration, and frequency of the perturbation. Then, lower species richness and abundance, shortening of the trophic webs, and the simplification of the trophic web are among the first observable damages contamination cause on microbial and protist communities [104]. Protists must at least tolerate the presence of the contaminant to achieve this function. Protists do not feed on hydrocarbons, but their grazing activity on the microorganisms that can keep the metabolization of the contaminant as high as another limiting factor allows them to.
Greater richness and abundance of ciliates species are associated with less perturbed areas; the greater the perturbation, the lesser species richness and abundance [105], regardless of the nature of the perturbing factor. For example, a significant reduction of ciliate diversity has been found in systems polluted by high hydrocarbon concentrations [106]. Medium concentrations only reduce the quantity of individuals from dominant species [106], while low concentrations produce an increase in the numbers of heterotrophic protists [107]. Saline accumulation forces the ciliates’ diversity to decrease as salinity values increase [108, 109]. In the same way, acidic pollution produces lower species richness and abundance as the environment becomes more acidic [110, 111], and the same pattern is observed with heavy metals’ contamination [104, 110].
Addition of organic matter in excess suddenly changes the base of production of the microbial food web, from phototrophs’ productivity to heterotrophic bacteria and yeasts’ productivity. The time of reaction is also different along the different microbial groups surviving the contamination event. Bacteria may start their biological activities several hours after the pollution event, whereas yeast and protists will delay from days to weeks, depending on the size of the organism.
Changes of primary producers from phototrophs to heterotrophs scale to functional groups, accommodating species richness and abundance of bacterivores protists, followed by omnivores. This is due to hydrocarbons stimulation of bacterial growth and the consequently increase of bacterivores species [112, 113]. Some species of genera
An increase in diversity and complexity of food webs are direct effects of these perturbations. Oil spill in deep waters increase the richness of the microbial community species and the complexity of their corresponding relationships, and the oil stimulated microbial activity supports greater variety of ciliates functioning along several trophic levels [117].
Other events of enriching oligotrophic systems with organic matter produce similar changes in the community structure of ciliates. Tirjaková and Vďačný [118] analyzed the changes in the communities of ciliates before and after a windstorm hit a stream, and they found a significant increase of ciliates’ species’ richness and abundance after the storm. Several weeks later, the community of ciliates presented the typical values of oligotrophic sites. The increase in resources availability is the factor indirectly responsible of these changes of ciliate community, but later, communities tend to return to states similar to the initial ones after resources exhaustion, which my take place around six months [118]. However, Shabarova et al. [119] report that the microbial community recovers from perturbation to a pre-flood state within two weeks after the event.
Regarding the connections’ shrinkage of the trophic networks, a gradual narrowing of the planktonic size spectrum has been reported in hypersaline lakes, correlated to salinity increases during the summer, resulting in a simplification of the community represented by the ciliated
Communities’ characteristic of hypersaline lakes are dominated by
Regarding the perturbances in the soil ciliated communities, similar effects have been described as in aquatic ecosystems. Exposure of ciliate communities to heavy metals induces a reduction in the biomass of ciliates and this effect lasts for 20 weeks [124]. Insecticides also generate a decrease in ciliates species immediately after contamination, they also generate a change in the dominance of ciliates, the bacterivores (
Protists in general, and ciliates in particular, play a key role in nutrient cycling and food web functioning in both aquatic and terrestrial ecosystems. In the world experiencing climate change and other kind of anthropogenic menaces, protists may be useful partners to tell us how aquatic and terrestrial systems are dealing with these issues while mesmerizing the observer with their great diversity of beautiful forms.
The authors declare no conflict of interest.
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