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.
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We 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!
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.
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Throughout 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\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\n
We 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:"7660",leadTitle:null,fullTitle:"Earthquakes - Impact, Community Vulnerability and Resilience",title:"Earthquakes",subtitle:"Impact, Community Vulnerability and Resilience",reviewType:"peer-reviewed",abstract:"This book is a collection of scientific papers on earthquake preparedness, vulnerability, resilience, and risk assessment. Using case studies from various countries, chapters cover topics ranging from early warning systems and risk perception to long-term effects of earthquakes on vulnerable communities and the science of seismology, among others. This volume is a valuable resource for researchers, students, non-governmental organizations, and key decision-makers involved in earthquake disaster management systems at national, regional, and local levels.",isbn:"978-1-78984-667-6",printIsbn:"978-1-78984-666-9",pdfIsbn:"978-1-78985-233-2",doi:"10.5772/intechopen.77465",price:119,priceEur:129,priceUsd:155,slug:"earthquakes-impact-community-vulnerability-and-resilience",numberOfPages:250,isOpenForSubmission:!1,isInWos:null,isInBkci:!1,hash:"4fa1f8e49ddf54c3caf69aa6100d2af8",bookSignature:"Jaime Santos-Reyes",publishedDate:"September 11th 2019",coverURL:"https://cdn.intechopen.com/books/images_new/7660.jpg",numberOfDownloads:12496,numberOfWosCitations:12,numberOfCrossrefCitations:15,numberOfCrossrefCitationsByBook:0,numberOfDimensionsCitations:25,numberOfDimensionsCitationsByBook:1,hasAltmetrics:0,numberOfTotalCitations:52,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"November 7th 2018",dateEndSecondStepPublish:"November 28th 2018",dateEndThirdStepPublish:"January 27th 2019",dateEndFourthStepPublish:"April 17th 2019",dateEndFifthStepPublish:"June 16th 2019",currentStepOfPublishingProcess:5,indexedIn:"1,2,3,4,5,6,7",editedByType:"Edited by",kuFlag:!1,featuredMarkup:null,editors:[{id:"188978",title:"Dr.",name:"Jaime",middleName:null,surname:"Santos-Reyes",slug:"jaime-santos-reyes",fullName:"Jaime Santos-Reyes",profilePictureURL:"https://mts.intechopen.com/storage/users/188978/images/system/188978.jpg",biography:"Jaime Santos-Reyes is a lecturer at the National Polytechnic Institute (IPN), Mexico, whose main research interests are safety management systems, technological & natural disaster risk, accident analysis, and systems' failure in general. He obtained a PhD from Heriot-Watt University, Edinburgh, Scotland, UK, in 2001. His internationally recognized papers have been listed for several years, within the TOP25 Hottest Articles. He (together with a multidisciplinary team) is conducting research on seismic risk and technological risk perception. He has created a research group called: Safety, Risk, Accident & Reliability Analysis - (SARACS) with the aim to conduct multidisciplinary research on system's failure in general.",institutionString:"National Polytechnic Institute",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"3",totalChapterViews:"0",totalEditedBooks:"1",institution:{name:"Instituto Politécnico Nacional",institutionURL:null,country:{name:"Mexico"}}}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,coeditorOne:null,coeditorTwo:null,coeditorThree:null,coeditorFour:null,coeditorFive:null,topics:[{id:"778",title:"Earthquake Engineering",slug:"engineering-environmental-engineering-earthquake-engineering"}],chapters:[{id:"67236",title:"Introductory Chapter: Earthquakes - Impact, Community Vulnerability, and Resilience",doi:"10.5772/intechopen.86284",slug:"introductory-chapter-earthquakes-impact-community-vulnerability-and-resilience",totalDownloads:732,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:null,signatures:"Jaime Santos-Reyes",downloadPdfUrl:"/chapter/pdf-download/67236",previewPdfUrl:"/chapter/pdf-preview/67236",authors:[{id:"188978",title:"Dr.",name:"Jaime",surname:"Santos-Reyes",slug:"jaime-santos-reyes",fullName:"Jaime Santos-Reyes"}],corrections:null},{id:"66486",title:"The IDEA Model as a Conceptual Framework for Designing Earthquake Early Warning (EEW) Messages Distributed via Mobile Phone Apps",doi:"10.5772/intechopen.85557",slug:"the-idea-model-as-a-conceptual-framework-for-designing-earthquake-early-warning-eew-messages-distrib",totalDownloads:868,totalCrossrefCites:3,totalDimensionsCites:6,hasAltmetrics:0,abstract:"Short response time available in the event of a major earthquake poses unique challenges for earthquake early warning (EEW). Mobile phone apps may be one way to deliver such messages effectively. In this two-phase study, several hundred participants were first randomly assigned to one of eight experimental conditions. Results of phase one afforded researchers the ability to reduce the number of conditions to four. Phase two consisted of five experimental conditions. In each condition, a 10 second EEW was delivered via a phone app. The four treatment conditions were designed according to elements of the IDEA model. The control condition was based on the actual ShakeAlert EEW computer program message being used by emergency managers across the US west coast at the time. Results of this experiment revealed that EEW messages designed according to the IDEA model were more effective in producing desired learning outcomes than the ShakeAlert control message. Thus, the IDEA model may provide an effective content framework for those choosing to develop such apps for EEW.",signatures:"Deanna D. Sellnow, Lucile M. Jones, Timothy L. Sellnow, Patric Spence, Derek R. Lane and Nigel Haarstad",downloadPdfUrl:"/chapter/pdf-download/66486",previewPdfUrl:"/chapter/pdf-preview/66486",authors:[{id:"222937",title:"Ph.D.",name:"Deanna",surname:"Sellnow",slug:"deanna-sellnow",fullName:"Deanna Sellnow"}],corrections:null},{id:"66621",title:"Knowledge and Perception on Seismic Risk of Students in Mexico City Before the 2017 Earthquakes",doi:"10.5772/intechopen.85556",slug:"knowledge-and-perception-on-seismic-risk-of-students-in-mexico-city-before-the-2017-earthquakes",totalDownloads:686,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"This paper presents some of the results of a cross-sectional study conducted in Mexico City in 2015–2016. The approach has been the application of a questionnaire to a sample size of n = 1489. Six high schools participated in the study that are located within the seismic zones of the city. Some of the results and conclusions are given below: (a) 95% of the students have experienced an earthquake and 71% considered that earthquakes cannot be predicted; however, 29% did not know this fact; (b) 82.2% of students were all aware of the likelihood of an earthquake occurrence sometime in the future. (c) One of the key conclusions is associated with the need to educate the residents of the capital city on a more realistic scale of the size of an earthquake; this could be the “Modified Mercalli Intensity Scale” or similar. (d) More generally, the residents of the city should be educated with urgency on these basic concepts. The more effective is the communication on risks and consequences, the better may be their preparedness to earthquakes.",signatures:"Tatiana Gouzeva, Diego Padilla-Pérez, Daniel Velázquez-Martínez, Vladimir Avalos-Bravo and José Lourdes Félix-Hernández",downloadPdfUrl:"/chapter/pdf-download/66621",previewPdfUrl:"/chapter/pdf-preview/66621",authors:[{id:"194713",title:"Dr.",name:"Vladimir",surname:"Avalos-Bravo",slug:"vladimir-avalos-bravo",fullName:"Vladimir Avalos-Bravo"},{id:"285299",title:"Dr.",name:"Tatiana",surname:"Gouzeva",slug:"tatiana-gouzeva",fullName:"Tatiana Gouzeva"},{id:"295821",title:"Dr.",name:"Diego",surname:"Padilla-Pérez",slug:"diego-padilla-perez",fullName:"Diego Padilla-Pérez"},{id:"295822",title:"Dr.",name:"Daniel",surname:"Velazquez-Marinez",slug:"daniel-velazquez-marinez",fullName:"Daniel Velazquez-Marinez"},{id:"295823",title:"Dr.",name:"Jose Lourdes",surname:"Félix-Hernández",slug:"jose-lourdes-felix-hernandez",fullName:"Jose Lourdes Félix-Hernández"}],corrections:null},{id:"66634",title:"Earthquake Disasters and the Long-Term Health of Rural Men in Chile: A Case Study for Psychosocial Intervention",doi:"10.5772/intechopen.84903",slug:"earthquake-disasters-and-the-long-term-health-of-rural-men-in-chile-a-case-study-for-psychosocial-in",totalDownloads:809,totalCrossrefCites:1,totalDimensionsCites:1,hasAltmetrics:0,abstract:"The article focuses on the long-term health of a rural male population exposed to a major earthquake event in Chile, in 2010. The results show that a majority of the male study participants considered that their physical and mental health had deteriorated over a 7-year span following the earthquake and that these impacts were strongest in men aged 65 years or more. In considering potential lessons for intervention, the results must be interpreted within the context of the construction of male identities in a rural community, informed by generally conservative values and binary male-female gender roles. The article concludes that health and social services workers and administrators providing interventions to male populations following earthquake must work to reduce the gap between the service offer and men’s real needs, which are frequently insufficiently understood and inadequately coded.",signatures:"Oscar Labra, Robin Wright, Danielle Maltais, Gilles Tremblay, Ray Bustinza and Gabriel Gingras-Lacroix",downloadPdfUrl:"/chapter/pdf-download/66634",previewPdfUrl:"/chapter/pdf-preview/66634",authors:[{id:"191776",title:"Dr.",name:"Oscar",surname:"Labra",slug:"oscar-labra",fullName:"Oscar Labra"}],corrections:null},{id:"67102",title:"Impacts of the 2015 Gorkha Earthquake: Lessons Learnt from Nepal",doi:"10.5772/intechopen.85322",slug:"impacts-of-the-2015-gorkha-earthquake-lessons-learnt-from-nepal",totalDownloads:2231,totalCrossrefCites:5,totalDimensionsCites:9,hasAltmetrics:0,abstract:"Nepal is highly vulnerable to a number of disasters for example: earthquakes, floods, landslides, fires, epidemics, avalanches, windstorms, hailstorms, lightning, glacier lake outburst floods, droughts and dangerous weather events. Among these disasters—earthquake is the most- scary and damaging. The effects of a disaster, whether natural or human induced, are often long lasting. The Gorkha earthquake of 25 April 2015 enormously affected human, socio-economic and other multiple sectors and left deep scars mainly in the economy, livelihood and infrastructure of the country. Besides the natural factors, the damages from disasters in Nepal are in increasing trend due to the human activities and inadequate proactive legislations. Fundamentally, the weak structures have been found as the major cause of damage in earthquakes. This underlines the need for strict compliance of building codes. Thus, proactive disaster management legislation focusing on disaster preparedness is necessary. This paper analyses and shows the critical gaps and responsible factors that would contribute towards seismic risk reduction to enable various stakeholders to enhance seismic safety in Nepal. Additionally, this chapter aims to pinpoint the deficiencies in disaster management system in Nepal with reference to the devastating Gorkha earthquake and suggest appropriate policy and advanced technical measures for improvement.",signatures:"Shiva Subedi and Meen Bahadur Poudyal Chhetri",downloadPdfUrl:"/chapter/pdf-download/67102",previewPdfUrl:"/chapter/pdf-preview/67102",authors:[{id:"285969",title:"Mr.",name:"Shiva",surname:"Subedi",slug:"shiva-subedi",fullName:"Shiva Subedi"},{id:"293220",title:"Dr.",name:"Meen",surname:"Paudyal Chhetri",slug:"meen-paudyal-chhetri",fullName:"Meen Paudyal Chhetri"}],corrections:null},{id:"66765",title:"Lifeline Interrelationships during the Tohoku Earthquake: Management of Disaster Analysis Reports Using Text Mining",doi:"10.5772/intechopen.84496",slug:"lifeline-interrelationships-during-the-tohoku-earthquake-management-of-disaster-analysis-reports-usi",totalDownloads:863,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"Although many studies have been undertaken in the area of lifeline interrelation, examinations based on the quantitative evaluation of post-earthquake lifeline interrelations have been limited. In this study, we present a new methodology to evaluate lifeline interactions, with an emphasis on the aspects of earthquake disasters. Terms related to other lifelines used in disaster reports are somewhat influenced by post-earthquake behaviors. In this study, the number of related terms was counted, and the relationships between the lifelines were quantitatively assessed for the 2011 Tohoku earthquake that occurred in Japan. The validation process included checks through academic reports as well as government reporting. We found that many lifelines were strongly dependent on the electric power systems, which gave no consideration to the accident at the Fukushima nuclear power plant. We confirmed further that the similar lifeline interrelation results could still be obtained regardless of the reporting used.",signatures:"Yasuko Kuwata",downloadPdfUrl:"/chapter/pdf-download/66765",previewPdfUrl:"/chapter/pdf-preview/66765",authors:[{id:"55698",title:"Dr.",name:"Yasuko",surname:"Kuwata",slug:"yasuko-kuwata",fullName:"Yasuko Kuwata"}],corrections:null},{id:"66324",title:"Resilience Science for a Resilient Society in Natural Disaster Prone Countries",doi:"10.5772/intechopen.85329",slug:"resilience-science-for-a-resilient-society-in-natural-disaster-prone-countries",totalDownloads:758,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"Recently, many destructive natural disasters occurred in the world. Therefore, the damage reductions and disaster mitigation for resilient society are very important and significant. For the implementation of these issues, we propose the resilience science including science, engineering, medicine, and social science. In social science, there are sociology, economics, psychology, law, pedagogy, etc. After 2011 earthquake in East Japan in which severe tsunami damages in a broad area occurred, the reconstruction and restoration activities in each area have been done; however, the progress speeds are not so rapid generally. One of reasons in which delayed reconstruction and restoration occurred is the shortage of pre-recovery plan and concept of future community in each area. In this chapter, we propose the resilience science for resilient society. The resilience science is based on multidisciplinary research fields, and the resilient society is defined as the society equipped with redundancy, robustness, elasticity, and safety. Especially, human resource cultivation is very important in resilience science for the resilient society. For the bright future, the resilience science for the resilient society based on human resource cultivation is indispensable.",signatures:"Yoshiyuki Kaneda",downloadPdfUrl:"/chapter/pdf-download/66324",previewPdfUrl:"/chapter/pdf-preview/66324",authors:[{id:"290117",title:"Dr.",name:"Yoshiyuki",surname:"Kaneda",slug:"yoshiyuki-kaneda",fullName:"Yoshiyuki Kaneda"}],corrections:null},{id:"66900",title:"Methodology for Community-Based Resilient Reconstruction",doi:"10.5772/intechopen.85790",slug:"methodology-for-community-based-resilient-reconstruction",totalDownloads:842,totalCrossrefCites:1,totalDimensionsCites:1,hasAltmetrics:0,abstract:"This chapter is to clarify the methodology of community-based resilient reconstruction based on 8 years of experiences after the Great East Japan Earthquake disaster occurred on March 11, 2011. Five stages had been clarified. The first stage is pairing support and grand design, which shows ideal perspectives on the area. It should be based on the scientific information and the historical and cultural accumulation of the area. It is better to prepare this stage before the disaster had occurred. The second stage is community-based workshop, for opening up refugees’ mind. It is essential to understand that the reconstruction should be carried out by refugees themselves. The third stage is decision-making process, together with refugees, local government, university, NPO, etc. How to create consensus building is the most critical issue. The fourth stage is the implementation. Supporting system should be created for the sustainability of community. The fifth stage is to develop the new community ties for the future generation and to return the benefits which they have received during the reconstruction.",signatures:"Mikiko Ishikawa",downloadPdfUrl:"/chapter/pdf-download/66900",previewPdfUrl:"/chapter/pdf-preview/66900",authors:[{id:"286814",title:"Prof.",name:"Mikiko",surname:"Ishikawa",slug:"mikiko-ishikawa",fullName:"Mikiko Ishikawa"}],corrections:null},{id:"68221",title:"Corporate Contributions to Community Resilience after the Great East Japan Earthquake Disaster",doi:"10.5772/intechopen.87947",slug:"corporate-contributions-to-community-resilience-after-the-great-east-japan-earthquake-disaster",totalDownloads:1029,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"In the Great East Japan Earthquake of March 11, 2011, municipalities in the northeastern coastal area of Japan suffered serious damage from the disaster. During such a huge disaster, it is difficult that the municipalities lead the regional recovery, and in order for the regional society to recover autonomously and efficiently from natural disasters, it is important that the communities in the devastated areas have the resilience. This paper focused on “corporate contribution” because the corporations have various resources that can support disaster recovery. Questionnaire surveys were conducted by mail to 1020 corporations including various industries and various sizes of corporations located in Iwanuma City and Natori City, which had been damaged by the Great East Japan Earthquake, analysis of memorial collection of sentences in which the emotions of disaster victims could be grasped, analysis of corporate network contribution, and analysis of corporate behavior based on the continuous interviews in the long term. Based on the results, I will give you a new concept “Gyo-Jyo (corporate contribution to community resilience)” and the information about what are the characteristics of corporate contributions to community resilience after the Great East Japan Earthquake disaster.",signatures:"Rui Fukumoto",downloadPdfUrl:"/chapter/pdf-download/68221",previewPdfUrl:"/chapter/pdf-preview/68221",authors:[{id:"286113",title:"Ph.D.",name:"Rui",surname:"Fukumoto",slug:"rui-fukumoto",fullName:"Rui Fukumoto"}],corrections:null},{id:"66155",title:"The Role of Placemaking as a Tool for Resilience: Case Studies from Post-Earthquake Christchurch, New Zealand",doi:"10.5772/intechopen.85119",slug:"the-role-of-placemaking-as-a-tool-for-resilience-case-studies-from-post-earthquake-christchurch-new-",totalDownloads:1220,totalCrossrefCites:1,totalDimensionsCites:3,hasAltmetrics:0,abstract:"In the aftermath of the 2010 and 2011 earthquakes in Christchurch, New Zealand, community-led temporary and adaptive urbanism filled a gap between the emergency response and recovery. In the space between response and recovery, the citizens of Christchurch showed their commitment to rethinking how they wanted to rebuild and then regenerate their city, leading to the embrace of collaborative processes, temporary and adaptive urbanism principles and a range of placemaking responses. In this chapter, the role of placemaking as a tool for post-disaster regeneration and resilience is considered by assessing three case study placemaking projects: the Re:START Mall, the Festival of Transitional Architecture (FESTA) and the placemaking programme at the Commons. Their development along with their success is considered within the context of the recovery of Christchurch and, in particular, how they align to the The Resilient Greater Christchurch Framework as it is set out in the Resilient Greater Christchurch Plan, in order to determine their role in building the resilience of Christchurch.",signatures:"Diane Brand, Hugh Nicholson and Natalie Allen",downloadPdfUrl:"/chapter/pdf-download/66155",previewPdfUrl:"/chapter/pdf-preview/66155",authors:[{id:"286899",title:"Prof.",name:"Diane",surname:"Brand",slug:"diane-brand",fullName:"Diane Brand"},{id:"286900",title:"Dr.",name:"Natalie",surname:"Allen",slug:"natalie-allen",fullName:"Natalie Allen"},{id:"286901",title:"Mr.",name:"Hugh",surname:"Nicholson",slug:"hugh-nicholson",fullName:"Hugh Nicholson"}],corrections:null},{id:"66014",title:"Multi-Hazard Assessment of Seismic and Scour Effects on Rural Bridges with Unknown Foundations",doi:"10.5772/intechopen.85158",slug:"multi-hazard-assessment-of-seismic-and-scour-effects-on-rural-bridges-with-unknown-foundations",totalDownloads:814,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"This chapter proposes a probabilistic framework for assessing seismic and scour effects on existing river-crossing bridge structures. The emphasis is on bridge structures in rural areas, for which it has been recognized that a large number of rural bridges have unknown foundation types and further are subject to both flooding-induced scour and seismic damage. With a review of the US-based rural bridges, this chapter presents a probabilistic framework for bridge performance assessment. Using a representative rural bridge model, the fragility results for the bridge reveal that scour tends to be beneficial in reducing structural damage at slight to moderate seismic intensities and to be detrimental in increasing collapse potential at high-level intensities. The demand hazard curves further quantify probabilistically the occurrence of local damage and global collapse, and systematically reveal the complex effects of scour as a hydraulic hazard on bridge structures.",signatures:"Xuan Guo, Zhaohua Dai and ZhiQiang Chen",downloadPdfUrl:"/chapter/pdf-download/66014",previewPdfUrl:"/chapter/pdf-preview/66014",authors:[{id:"220361",title:"Prof.",name:"ZhiQiang",surname:"Chen",slug:"zhiqiang-chen",fullName:"ZhiQiang Chen"},{id:"284942",title:"Dr.",name:"Zhaohua",surname:"Dai",slug:"zhaohua-dai",fullName:"Zhaohua Dai"},{id:"285217",title:"Dr.",name:"Xuan",surname:"Guo",slug:"xuan-guo",fullName:"Xuan Guo"}],corrections:null},{id:"66024",title:"Earthquake-Generated Landslides and Tsunamis",doi:"10.5772/intechopen.84807",slug:"earthquake-generated-landslides-and-tsunamis",totalDownloads:812,totalCrossrefCites:3,totalDimensionsCites:4,hasAltmetrics:0,abstract:"Large earthquakes generate tsunamis, but when it also triggers a landslide, the tsunami may become enormous. Slide scars on the continental shelf of the North Atlantic Ocean show this. For estimating the tsunami, a translatory wave theory has been suggested. Slide data are used to estimate the amplitude of the displacement wave. The amplitudes are used to obtain wave heights at a reference point outside the breaker zone. Energy transmission formulas are used to find the wave height transfer coefficients from the source area to a reference point. Tsunami risk from several sources at a reference point is quantified using stochastic processes, and estimations of a hazard curve for the probability of landslide occurrence are carried out. The sensitivity of the hazard curve to uncertainties in determining the wave height from the individual sources turns can be evaluated. In two case studies, the Tohoku tsunami and earthquake in 2011 in Japan is found to be caused by a coseismic slip and a landslide in combination, and a hazard curve for a reference point south of Iceland is found for tsunamis in the North Atlantic Ocean.",signatures:"Jonas Eliasson",downloadPdfUrl:"/chapter/pdf-download/66024",previewPdfUrl:"/chapter/pdf-preview/66024",authors:[{id:"284319",title:"Dr.",name:"Jónas",surname:"Elíasson",slug:"jonas-eliasson",fullName:"Jónas Elíasson"}],corrections:null},{id:"65945",title:"Kinematics of Slow-Slip Events",doi:"10.5772/intechopen.84904",slug:"kinematics-of-slow-slip-events",totalDownloads:833,totalCrossrefCites:1,totalDimensionsCites:1,hasAltmetrics:0,abstract:"Large earthquakes are often preceded or followed by slow-slip events, which when better understood may help better understand the mechanisms of the earthquakes and the possibility of their prediction. This chapter summarizes kinematic values of large slow-slip events observed in Circum-Pacific subduction zones and creep events observed along strike-slip faults in California. The kinematic parameters include maximum slip S, duration T, rupture length L, rupture width, magnitude M, slip velocity VS, rupture velocity VR, and maximum slip/rupture length ratio S/L. For a large surface and subsurface creep event in California: S = 0.9–2.5 cm, T = 2–5 day, L = 6–8 km, W = 3–4 km, M = 4.7–4.8, VS = 0.4–0.5 cm/d, VR = 1.6–3.0 km/d, and S/L = 0.3–1.5 × 10−6. For a large short-term slow-slip event in Circum-Pacific subduction zones: S = 1–20 cm, T = 2–50 day, L = 20–260 km, W = 10–90 km, M = 5.6–7.0, VS = 0.1–0.8 cm/d, VR = 2–20 km/d, and S/L = 0.3–1.5 × 10−6. The latter kind of events have larger sizes in slip, duration, rupture length/width, and magnitude than the former, but are comparable in slip velocity, rupture velocity, and S/L ratio. The kinematic behaviors of both are similar, despite their large difference in temperature, pressure, and composition of the fault-zone materials. The larger size of the latter is probably due to their larger inertia caused by their larger overburden. Compared with normal earthquakes, the slip and rupture velocities of both are smaller by many orders of magnitude. But their S/L values, and thus stress drops, are smaller by only one or two orders of magnitude. For a large long-term slow-slip event in the subduction zones: S = 1–50 cm, T = 50–2500 day, L = 40–1000 km, W = 30–750 km, M = 6.0–7.7, VS = 0.01–0.10 cm/d, VR = 0.1–2 km/d, and S/L = 0.1–2 × 10−6. The estimated slip, duration, rupture length, and magnitude values are larger than the short-term events, but the average slip and rupture velocities are much smaller. 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1. Introduction
An amoeba is very interesting organism to study because it grows as single cells and develops as multi‐cellular organisms. They present a range of developmental processes which can be used to study of any molecular pathways like glycolysis or gluconeogenesis pathways [1]. During evolution, the amoebozoa generated a large number of species which goes through the similar developmental stages, from unicellular to multi‐cellular stages [1–4]. The well‐characterized amoebozoan species is Dictyostelium discoideum, which is easy to study as compared to mammalian cell [5–7]. D. discoideum uses many similar signals and contains similar pathways that are presented in plants and animals [1]. D. discoideum or cellular slime mold is simple eukaryotic microorganism which generally grows in forest soil and decaying leaves. This amoeba feeds on bacteria and grows as single cells [2, 3]. The development of D. discoideum is simpler than that of mammalian cells.
2. The gluconeogenesis process in eukaryotic cell
Gluconeogenesis is a process by which carbohydrate is synthesized from non‐carbohydrate precursors like oxaloacetate and pyruvate (Figure 1). In the first step of the gluconeogenesis process, oxaloacetic acid is synthesized from pyruvic acid. On the other hand, in the citric acid cycle, oxaloacetic acid reacts with acetyl‐CoA. So, at low concentration of acetyl‐CoA and high concentration of ATP, gluconeogenesis proceeds. Gluconeogenesis starts in the mitochondria of the cells. In the first step, carboxylation of pyruvate occurs by pyruvate carboxylase enzyme and it forms oxaloacetate by using one ATP molecule. Oxaloacetate is reduced to malate by using NADH. After this step, the remaining steps of gluconeogenesis process occur in the cytosol. In the next step, malate is oxidized to oxaloacetate using NAD+. Oxaloacetate is first decarboxylated, and after that, it is phosphorylated by using the enzyme, PEP carboxykinase, and one GTP. In the next step, PEP converted into 2‐phosphoglycerate, 3‐phosphoglycerate and then 1,3‐bisphosphoglycerate by the enzyme enolase, phosphoglycerate mutase and phosphoglycerate kinase, respectively. In the next step of this reaction, 1,3‐bisphosphoglycerate converts into glyceraldehyde 3‐phosphate by the enzyme glyceraldehyde phosphate dehydrogenase. Now, the glyceraldehyde 3‐phosphate converts into fructose 1,6‐bisphosphate via two ways: one is direct conversion and another through the intermediate component called dihydroxyacetone phosphate. In the next step, fructose 1,6‐bisphosphate converts into fructose 6‐phosphate, using an enzyme, fructose 1,6‐bisphosphatase, one water molecule, and releasing one phosphate. This step is the rate‐limiting step in gluconeogenesis process. Glucose‐6‐phosphate is formed from fructose 6‐phosphate followed by glucose by the enzyme glucose‐6‐bisphosphatase. The reaction of the glucose formation occurs inside the endoplasmic reticulum, specifically in the lumen, where glucose‐6‐phosphate is hydrolyzed and produces glucose and releases an inorganic phosphate [8]. 3.The developmental stages of Dictyostelium discoideum.
Figure 1.
Gluconeogenesis process in the eukaryotic cell.
About 80 years ago, Ken Raper isolated D. discoideum from the forest floor at North Carolina [9]. He observed that when the cells had a depletion of food, they aggregated into mounds [9]. John Bonner showed that when the cells were in the starving condition, they secreted a chemical which acts like chemoattractant and the cells responded by moving up the gradient [10]. After 20 years, Konijn et al. showed that the chemoattractant was cAMP [11]. After this discovery, D. discoideum considered as a model organism to study chemotaxis and developmental biology. The connection between cell signaling pathways and biochemical pathways was established by using this organism. The pathways for cAMP synthesize, the surface receptors for cAMP and many other cell signaling to biochemical and molecular biological techniques were established by using D. discoideum [12–15]. The developmental stages of D. discoideum started from slug‐shaped structures and go till the formation of the fruiting body [1, 2, 16]. Pre‐spore and pre‐stalk cells at the slug stage formed spores and stalk cells in fruiting bodies, and it was also found that pre‐stalk cells were at the front of the slugs, and pre‐spore cells were all in the back [2, 16]. There were 20‐fold differences between the size of slugs and the total number of individual cells in each slug. This variation showed that there was some intracellular signal which determines the proportions of pre‐spore and pre‐stalk cells. Soderbom and Loomis showed that the pre‐spore cells synthesize an inhibitor which inhibits pre‐spore differentiation, and the pre‐spore cells were resistant to this inhibitor [2, 17, 18]. This mechanism was responsible for size invariance of the slugs [18]. The amoeba went through three cycles when it was facing starvation [7].
3.1. The microcyst
Encystment was a very common process to amoebae, but it was not known for D. discoideum. In microcyst stage, each cell elaborated into a two layered cellulose coat and went to the dormant stage [7].
3.2. The macrocyst
In this stage of the sexual cycle, cells of two mating types fused [2]. Under wet condition, the macrocyst form, which had three layered cellulose coat at maturity. After fusion, the cells formed giant cells which had at least two nuclei or many nuclei. This fused structure attracted other amoebae by chemotaxis to cAMP. The endocytes were formed by engulfing these cells, and after that the giant cells produced meiotic offspring [7]. Macrocysts were formed from endocytes including hundred of cells.
3.3. Fruiting bodies
The fruiting body was formed through complex and polarized cell movements. In this stage, cells were not engulfed to form endocytes because one cell was recognized by the other cell. For the formation of fruiting body of D. discoideum, cells did chemotaxis and cells were more elaborated and involved a relay mechanism. But this mechanism either suppressed or did not exist during the macrocyst formation. Fruiting body formed by aggregation of one lakh cells. In this case, cells were adhesive in nature and moved among each other and able to distinguish between cAMP and other molecules [7].
During mid‐developmental stage, D. discoideum can choose between two different pathways, one is from the finger stage, it can directly precede to culmination, or it can fall over to form a phototactic, migratory slug. This migratory stage is important for cells to find an appropriate site for fruiting body formation [19]. Slugs prefer dark and low ionic strength environment [20]. Two cell types were there inside the slug which implied that they were connected with the signaling system [7]. Twenty percentage cells died for the formation of the stalk of the fruiting body, and eighty percentage cells survive by the formation of spores. Figure 2 represents the developmental stages of D. discoideum.
Figure 2.
Lifecycle of Dictyostelium discoideum.
4. Size of the aggregates of D. discoideum is depending upon the gluconeogenesis pathway indirectly
The group size of D. discoideum is regulated by a negative feedback pathway mediated by counting factor (CF) which consists of at least five protein complexes [24]. During the early developmental stage, the counting factor (CF) breaking up the big aggregate of cells to the smaller aggregates of about 2 × 104 cells [21]. High levels of CF decrease cell‐cell adhesion and also decrease the amplitude of cAMP and increase random motility. The glucose metabolism is affected by the CF, and it decreases the CF glucose levels [22]. CF decreases the activity of the gluconeogenic enzyme, glucose‐6‐phosphatase, which decreases the level of glucose in the cell with the high secretion of CF [23, 24]. In that case, if glucose has been added externally then the size of the fruiting bodies get increases [24]. This process alters the intermediates of the metabolic pathways, such as pyruvate and lactate [22]. Jang and Gomer showed that, if the cells exposed to CF, the CF has very small effect on amylase or glycogen phosphorylase, enzymes involved in glucose production from glycogen [24]. On the other hand, it has a huge effect on glycolysis pathway. If the CF is high, then it is inhibited the glucokinase activity, but it does not regulate phosphofructokinase (enzyme responsible for glycolysis pathways). CF showed some effect upon the enzyme involved in a gluconeogenesis pathway such as fructose‐1,6‐bisphosphatase and glucose‐6‐phosphatase. The fructose‐1,6‐bisphosphatase is not regulated by CF, whereas glucose‐6‐phosphatase is regulated by CF [24].
The size of the terminal structures of D. discoideum is depending upon the cell surrounded by the number of cells, so the initiation of development is not started with the significant growth of the cell. The secretion of a protein complex is very important to control the size of the aggregates. Large aggregates initially have higher CF levels which can modify gluconeogenesis, and the other metabolic pathways alters the level of metabolites [1, 24]. After that, the level of CF drops down which increases cell‐cell adhesion and because of that the random motility decreases which stabilize the smaller aggregates.
5. Gluconeogenesis process affected during the differentiation of myxamoebae of the cellular slime mold
Myxamoeba is a naked amoeboid uni‐nucleate protoplast that lacks both cilia and flagella. In the life cycle of D. discoideum, they had gone through the vegetative state, differentiation state as independent amoeboid cells which called as myxamoebae [25]. If the myxamoebae was grown in different media, with varying carbohydrate content [26], then the chemical composition [27], enzyme composition [28], and physiological behavior [29] got changed inside the cell. After that, if these cells put in the moist condition, the cells formed slug [5, 25]. Carbohydrate content changed inside the D. discoideum at different stages of development [30]. So the gluconeogenesis process alters during the carbohydrate conversion process, the energy was coming from cellular protein, RNA, and dry weight to complete this process [25, 30–33].
Wright et al. showed in their kinetic model, the glycogen content of the cell remained constant during the early developmental stage but it decreased when the culmination process occurred [34]. White and Sussman suggested that the glycogen content of axenic cells was small [30]. They also showed that the glycogen initially decreased because of the consumption of the bacterial glycogen for the development of axenically grown myxamoebae. Wright et al. assume in their model, during differentiation of D. discoideum, there was no gluconeogenesis process occurs [34]. Cleland and Coe showed some evidence of the presence of gluconeogenesis process during the differentiation of myxamoebae [31]. However, Hames et al. suggested that initially, cells had low glycogen but during differentiation, the glycogen content increased significantly which depicts that the gluconeogenesis process had a huge effect on the differentiation mechanism of the cell [25, 35].
In the absence of glucose and the presence of very low concentrations of glycogen, myxamoebae grow and degrade all the glycogen during 4 h of development [35]. Glycogen is synthesized during the late developmental stages (5–15 h) and finally broken down by the cell to synthesize saccharide. Hames and Ashworth showed that the amount of glycogen synthesized during the late developmental stage is larger than the glycogen content of the vegetative cells [25]. This glycogen synthesis occurs during gluconeogenesis process when the cellular glucose remains at a constant low concentration. During differentiation, myxamoebal glycogen is not stored, but the gluconeogenesis process still can occur, if the cells initially have a large amount of glycogen.
6. Discussion
D. discoideum is a well‐characterized eukaryotic system which grows as single cells and develops as multi‐cellular organisms. The development of D. discoideum is simpler than that of mammalian cells. They use many similar signals and contain similar pathways that are presented in plants and animals so it is the best biological system to study the molecular pathways like glycolysis or gluconeogenesis pathways, which can correlated with the mammalian system. D. discoideum considered as a model organism to study chemotaxis and developmental biology. D. discoideum obtained energy through lysosomal degradation and phagocytosis process [36]. Because of that intra‐lysosomal nutrient levels are increasing so, the lysosome and the vacuole collect amino acids and other nutrients from cellular components through the autophagy process [37]. This process has been correlated with the gluconeogenesis process. The counting factor (CF) or the protein complex regulates the size of the aggregates of D. discoideum, which is indirectly affecting the gluconeogenesis pathway. CF showed some effect upon the enzyme involved in a gluconeogenesis pathway such that fructose‐1,6‐bisphosphatase and glucose‐6‐phosphatase. The fructose‐1,6‐bisphosphatase is not regulated by CF, whereas glucose‐6‐phosphatase is regulated by CF, which acts as one of the size regulating factor for the aggregates of D. discoideum. Large aggregates initially have higher CF levels which can modify gluconeogenesis, and the other metabolic pathways, and also alters the level of metabolites [1, 24]. After that, the level of CF drops down which increases cell‐cell adhesion and because of that the random motility decreases which stabilize the smaller aggregates. The glycogen synthesis during the developmental process of D. discoideum causes by the gluconeogenesis pathway. During differentiation, myxamoebal glycogen is not stored but the gluconeogenesis process still can occur, if the cells initially have a large amount of glycogen. So gluconeogenesis pathway is very important for the development of the cells, this is one of the best eukaryotic system in which the process can be studied.
\n',keywords:"gluconeogenesis, eukaryotic system, Dictyostelium discoideum",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/53886.pdf",chapterXML:"https://mts.intechopen.com/source/xml/53886.xml",downloadPdfUrl:"/chapter/pdf-download/53886",previewPdfUrl:"/chapter/pdf-preview/53886",totalDownloads:1479,totalViews:510,totalCrossrefCites:0,totalDimensionsCites:0,totalAltmetricsMentions:0,impactScore:0,impactScorePercentile:19,impactScoreQuartile:1,hasAltmetrics:0,dateSubmitted:"June 22nd 2016",dateReviewed:"December 12th 2016",datePrePublished:null,datePublished:"July 19th 2017",dateFinished:"January 17th 2017",readingETA:"0",abstract:"Dictyostelium discoideum or cellular slime mold is simple eukaryotic microorganism, which generally grows in forest soil and decaying leaves. This amoeba feeds on bacteria and grows as single cells. The development of Dictyostelium discoideum is simpler than that of mammalian cells. It uses many of the same signals that are found to function in higher eukaryotic organisms like plants and animals. Dictyostelium discoideum is an excellent system in which to study metabolic pathways which are simpler than that of the complex systems like mammalian system. Glucose is metabolized in glycolysis to yield pyruvate and lactate and further metabolized in the tricarboxylic acid cycle. Glucose can be polymerized into glycogen in addition to glycolysis process. In a metabolic pathway, the generation of glucose from certain non‐carbohydrate carbon substrates is called gluconeogenesis. In Dictyostelium discoideum, glucose is synthesized by the breakdown of pyruvate. Glycogen phosphorylase and amylase break down glycogen to form glucose. Glycogen synthase and glycogen phosphorylase are the key enzymes for the regulation. Both the enzyme equally regulated the process simultaneously, so that when one is activated, the other is deactivated. During gluconeogenesis, glucose is synthesized from pyruvate but sometimes during this process, three enzymes, glucose‐6‐phophatase, fructose‐1,6‐bisphosphatase, and phosphoenolpyruvate carboxykinase catalyze an irreversible reaction.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/53886",risUrl:"/chapter/ris/53886",book:{id:"5614",slug:"gluconeogenesis"},signatures:"Richa Karmakar",authors:[{id:"193984",title:"Dr.",name:"Richa",middleName:null,surname:"Karmakar",fullName:"Richa Karmakar",slug:"richa-karmakar",email:"richa1503@gmail.com",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",institution:{name:"University of California, San Diego",institutionURL:null,country:{name:"United States of America"}}}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. The gluconeogenesis process in eukaryotic cell",level:"1"},{id:"sec_2_2",title:"3.1. The microcyst",level:"2"},{id:"sec_3_2",title:"3.2. The macrocyst",level:"2"},{id:"sec_4_2",title:"3.3. Fruiting bodies",level:"2"},{id:"sec_6",title:"4. Size of the aggregates of D. discoideum is depending upon the gluconeogenesis pathway indirectly",level:"1"},{id:"sec_7",title:"5. Gluconeogenesis process affected during the differentiation of myxamoebae of the cellular slime mold",level:"1"},{id:"sec_8",title:"6. Discussion",level:"1"}],chapterReferences:[{id:"B1",body:'Loomis, W. F. (2014). Cell signaling during development of Dictyostelium. Dev. Biol.; 391:1-16.'},{id:"B2",body:'Raper, K. B. (1984). The Dictyostelids. Princeton University Press, Princeton, NJ.'},{id:"B3",body:'Eichinger, L., Pachebat, J. A., Glockner, G., Rajandream, M. A., Sucgang, R., Berriman, R., Song, J., Olsen, R., Szafranski, K., Xu, Q., Tunggal, B., Kummerfeld, S., Madera, M., Konfortov, B. A., Rivero, F., Bankier, A. T., et al. (2005). The genome of the social amoeba Dictyostelium discoideum. Nature; 435:43-57.'},{id:"B4",body:'Schilde, C., Schaap, P. (2013). The amoebozoa. Methods Mol. Biol.; 983:1-15.'},{id:"B5",body:'Bonner, J. T. (1959). The Cellular Slime Molds. Princeton University Press, Princeton, NJ.'},{id:"B6",body:'Loomis, W. F. (1975). Dictyostelium discoideum—A Developmental System. Academic Press, New York.'},{id:"B7",body:'Kessin, R. H. (2001). Dictyostelium—Evolution, Cell Biology, and the Development of Multicellularity. Cambridge University Press, Cambridge, UK.'},{id:"B8",body:'Nelson, D. L., Cox, M. M. (2000). Lehninger Principles of Biochemistry. Worth Publishers, USA.'},{id:"B9",body:'Raper, K. B. (1935). Dictyostelium discoideum, a new species of slime mold from decaying forest leaves. J. Agric. Res.; 50:135-147.'},{id:"B10",body:'Bonner, J. T. (1947). Evidence for the formation of cell aggregates by chemotaxis in the development of the slime mold Dictyostelium discoideum. J. Exp. Zool.; 106:1-26.'},{id:"B11",body:'Konijn, T. M., VandeMeene, J. G. C., Bonner, J. T., Barkley, D. S. (1967). 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Cell.; 4:72-81.'},{id:"B25",body:'Hames, B. D., Ashworth, J. M. (1974). The metabolism of macromolecules during the differentiation of myxamoebae of the cellular slime mould Dictyostelium discoideum. Biochem. J.; 142:301-315.'},{id:"B26",body:'Watts, D. J., Ashworth, J. M. (1970). Growth of myxamoebae of the cellular slime mould Dictyostelium discoideum in axenic culture. Biochem. J.; 119:171-174.'},{id:"B27",body:'Weeks, G., Ashworth, J. M. (1972). Glycogen Synthetase and the control of glycogen synthesis in the cellular slime mould Dictyostelium discoideum during the growth (Myxamoebal) phase. Biochem. J.; 126:617-626.'},{id:"B28",body:'Ashworth, J. M., Quance, J. (1972). Enzyme synthesis in myxamoebae of the cellular slime mould Dictyostelium discoideum during growth in axenic culture. Biochem. J.; 126:601-608.'},{id:"B29",body:'Ashworth, J. M., Watts, D. J. (1970). Metabolism of the cellular slime mould Dictyostelium discoideum grown in axenic culture. Biochem. 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D., Weeks, G., Ashworth, J. M. (1972). Glycogen synthetase and the control of glycogen synthesis in the cellular slime mould Dictyostelium discoideum during cell differentiation. Biochem. J.; 126:627-633.'},{id:"B36",body:'Neuhaus, E. M., Almers, W., Soldati, T. (2002). Morphology and dynamics of the endocytic pathway in Dictyostelium discoideum. Mol. Biol. Cell.; 13:1390-1407.'},{id:"B37",body:'Efeyan, A., Comb, W. C., Sabatini, D. M. (2015). Nutrient‐sensing mechanisms and pathways. Nature; 517:302-310.'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Richa Karmakar",address:"richa1503@gmail.com",affiliation:'
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1. Introduction
Plants constitute vast and diverse niches for endophytic organisms, and there is not a single plant species devoid of them. The most up-to-date definition for endophytes defines them as the microorganisms isolated from surface-sterilized plant tissues, which do not cause any noticeable harm to their host plants [1, 2]. The most abundant and common microbes living as endophytes are bacteria and fungi [3]. Endophytic bacteria are present in any kind of plant, from ferns and bryophytes to mono and dicotyledonous species [4]. In nature, mainly the intercellular spaces of the plant host are colonized by the endophytic bacteria [1, 5, 6]. But, endophytes have been also found in intracellular spaces of grapevine, barley, tobacco, Arabidopsis, and pine [7], suggesting that legumes may also have intracellular endophytes.
The endophytic bacterial communities make significant contributions to growth promotion and plant health in mutualistic (even symbiotic) relationships. The plant host protects the bacteria from the environment, while the endophytic community provides several benefits to the host. The benefits for the plant may include nutrient assimilation (such as nitrogen, phosphorus, or iron), growth stimulation, defense against pathogens, and/or protection against environmental stresses [8, 9]. Some of these effects might be altered when the plant is under stress [10].
The use of these natural symbionts/mutualists offers an opportunity to maximize legume crop productivity while reducing the environmental impacts of agriculture. For decades, most of the studies (and agricultural applications) have been about the effects of individual strains of bacteria, but recently with the bloom in bioinformatics and sequencing technology development, the knowledge about the plant microbiota has burst, and the potential to use and manipulate complex bacterial communities has started to be the target of a large research community.
2. Plant endophytic microbiome
In natural environments, the intracellular spaces of legumes are inhabited by numerous microorganisms, such as virus, fungi, nematodes, and bacteria. Here we focus on bacterial endophytes that benefit the plant in some way. Those bacteria colonize the host by several mechanisms, such as natural opening or injures and proliferate within the host. There is a huge taxonomic and functional diversity of endophytic bacteria, adapted to the microenvironments that the plant host provides. That diversity will be shaped by the microbial community members, the plant host, and the environmental conditions.
2.1 Colonization and distribution within the host plant
Colonization mechanisms vary with the type of interaction between the host and the bacteria and the life cycle of the microbe. Overall, most of the endophytic bacteria enter the plant through the roots. Since the microbial diversity decreases from the root to the leaves, it has been proposed that most of the microbes colonize the plant through the roots and proliferate to aboveground tissues [11] (Figure 1). Endophytic bacteria are usually “recruited” by plant host root exudates, such as organic acids, amino acids, and proteins [12, 13]. Once the bacteria are close to the root surface, they enter through lateral root emergence areas or other openings, caused by wounds or mechanical injuries. In the early stages, most of the endophytes are first observed in root hairs and subsequently in the root cortex [14]. However, endophytes can also colonize the leaves through the stomata, injuries in the epidermis, or introduced by vectors. In leaves, bacterial endophytes have been observed in the intercellular spaces of mesophyll, substomatal areas, and xylem tissues [15, 16].
Figure 1.
Diversity gradient of bacterial endophytic microbiota and growth promotion mechanisms to legumes. Legumes are surrounded and interact with bacteria in the soil and air (epiphytic bacteria in the rhizosphere and phyllosphere) and in the inter- and intra-cellular spaces (endosphere). Those bacteria can be saprophytic, pathogenic, or beneficial for the plant. The beneficial bacteria can promote plant growth by direct and indirect mechanisms. Direct mechanisms include phytohormone, volatiles, and other compounds production and facilitation of nutrient assimilation. Indirect mechanisms include pathogen and abiotic stress protection. ISR, induced systemic resistance.
In addition, the habit of the microbe conditions its colonization strategy. For example, obligate endophytes, which depend on the plant metabolic activity for their survival, are usually transmitted to the seed (vertical transmission) and spread inside the plant or through the action of a vector. On the contrary, most of the facultative endophytes, which have a free life in the soil and colonize the plant during some stage of their life cycle, colonize the plant through occasional wounds [17].
The colonization process itself alters host plant physiology (in a process called “niche construction” from the microbe’s point of view) by defense alterations or direct shift of the host metabolism [18]. Those microenvironment changes can affect the local microbiome structure and functions, by altering relationships among bacterial species and within the host. Furthermore, under particular conditions, part of the response of the plant will stimulate or recruit specific endophytes, which may contribute to survival or tolerance of that condition [19, 20]. It was proved in tomato cultivars that the transplant of the rhizosphere from a resistant to a susceptible cultivar suppressed Ralstonia solanacearum disease symptoms. They found a highly abundant flavobacterial genome in the resistant cultivar rhizosphere, and the isolated flavobacteria suppressed disease symptoms in the susceptible cultivar in pots [21]. In legumes, it was reported that Fusarium-resistant common bean cultivars showed a higher abundance of Pseudomonadaceae, Bacillaceae, Solibacteraceae, and Cytophagaceae families [22], but no further inquiries have been reported.
Another aspect affecting the colonization process of the endophytic bacteria is the host defenses. Endophytes live in the same environment as many plant pathogens and share close similarities with them. Microbe- or pathogen-associated molecular patterns (MAMPs/PAMPs) are conserved and necessary for microbial survival, but plants have evolved multiple receptors to recognize them and induce the plant immune system. Then, the colonization of endophytic bacteria triggers plant defenses, and the process needs to be avoided or blocked by the beneficial endophytes to be able to colonize and proliferate within the host [2, 23, 24]. It is not well understood yet how the beneficial bacteria overcome the defenses, but a few mechanisms have been unraveled, including the blockage of MAMPs and defense signaling [25]. The beneficial bacteria Bacillus subtilis avoid a strong defensive response in the host by blocking the detection of their own flagellin by the secretion of the flagellin-binding peptide subtilomycin [25, 26]. Another mechanism is the secretion of bacterial antioxidant enzymes, such as superoxide dismutases and glutathione-S-transferases to detoxify the reactive oxygen species that signals the plant defense [27]. An alternative mechanism is the suppression of salicylic acid (SA)-mediated defense signaling. Sinorhizobium fredii HH103 with defective type III secretion system (T3SS) is unable to suppress SA-dependent defenses and subsequently fails to promote nodulation on the host [28], indicating that the suppression of the SA-dependent defense is critical for endophyte colonization. Some of those mechanisms have not been reported in legumes, but if those bacteria are colonizing legumes, similar mechanisms might be in action.
The establishment of the endophytic bacterial community in the legume host is a complex and dynamic process that has been studied mostly in fragments and simplified systems (usually one bacterial strain in one host under one or a few conditions), and it must be further understood to take the best advantages of their potential benefits for legume agriculture.
2.2 Endophytic bacterial diversity
There is an enormous diversity of bacterial endophytes in legumes, considering that the rhizobia are also endophytes. The interaction of rhizobia and legumes has been studied for more than a century [29]. Since then, many rhizobial endophytic bacteria were isolated from different legumes, particularly root and nodule tissue. These bacteria can establish a symbiotic interaction, induce the formation of new organs in roots and stems called nodules, and fix atmospheric nitrogen. In addition, the so-called “new rhizobia” (or noncanonical rhizobial genera) of Alfa- and Beta-Proteobacteria has been reported in the last decades. They can form nodules and fix nitrogen and mainly belong to Microvirga spp. and Burkholderia spp., respectively [30]. Other non-nitrogen-fixer endophytes are present in nodules and sometimes improve nodule formation [31, 32, 33]. For instance, Hoque et al. [34] isolated rhizobia and non-rhizobia endophytes from two wild Acacia species from Australia, and nodules were produced by species of the genera Rhizobium, Ensifer, Mesorhizobium, Burkholderia, Phyllobacterium, and Devosia, much more than expected. In addition, rhizobial species were isolated from other plant tissues apart from nodules [3].
Overall, from a large number of bacterial genera present in legumes, the most frequent ones (excluding rhizobia) are Agrobacterium, Bacillus, Enterobacter, and Pseudomonas, followed by Acinetobacter, Arthrobacter, Curtobacterium, Devosia, Dyella, Herbaspirillum, Klebsiella Micromonospora, Microbacterium, Mycobacterium, Ochrobactrum, Paenibacillus, Pantoea, Rhodopseudomonas, Serratia, Staphylococcus, and Sphingomonas ([3, 9, 21], and reference therein) (Tables 1 and 2).
Rhizobium phaseoli, Bacillus tequilensis, B. altitidinus, B. tequilensis, B. siamensis, B. subtilis, Pantoea dispersa, Paenibacillus illinoisensis, Kosakonia oryzendophytica, Rhizobium mayense, P. dispersa
IAA; ACC-DA; P, Zn, and Si solubilization, siderophore
Culture-dependent studies of the endophytic bacterial microbiota in legume crops.
ACC, 1-aminocyclopropane-1-carboxylate; ACC-DA, ACC deaminase activity; IAA, indole-acetic acid; BNF, biological nitrogen fixation; Develop., developmental stages; MG, meta-genomics; MT, meta-transcriptomics; N.D. not determined; N.S., not significant; Morph & Bioch., morphological and biochemical characterization, Treat, treatment or factor affecting microbiome.
2.3 Factors affecting diversity
The composition, diversity, and abundance of the endophytic microbiome are influenced by the soil microbial pool; the plant host identity and status (genotype, development, and physiology); agricultural practices; and climate and environmental conditions (such as temperature, water supply, and nutrients) [8, 16, 71]. Comparisons among plant species (canola, wheat, pea, and lentil) in different locations and soil types pointed to the genotype influence as the highest effect determining endophyte diversity ([72] in Table 1). However, when considering close Medicago genotypes (intraspecies comparison), the host genotype effect was not significant (1% of contribution to the total variance), but both soil and plant genotypes were significant for the root microbiota diversity [53]. In the case of the leaf microbiome, the soil reduces its relative importance, since some bacteria colonize it from underground organs, but others enter through stomata or vectors [46]. Broadly, the soil limits the available microbial pool, while the host genotype is a relevant barrier for colonization. Agricultural practices could directly affect the microbiome by chemical applications or through changes in the host physiology. The effects of biotic and abiotic factors shaping the endophytic bacteria communities in plants were reviewed by Papik et al. [73]. In addition, the actual diversity could be masked by the method used to describe it (such as culture-dependent or -independent, see Section 2.4) [16].
2.4 How to study microbiome diversity
Natural communities of endophytic bacteria are conventionally studied using culture-dependent and -independent methods [74]. Culture-dependent methods imply the extraction of the microbes and their growth in synthetic media. Those strategies allow to isolate the microbe and further study them in vitro and in manipulative experiments, but they strongly underestimate the number of bacteria (and the diversity of the community), as cultivable bacteria usually represent only 0.001–1% of the actual bacteria in a sample [16, 75]. Recently, Hartman et al. [52] isolated 200 bacteria strains that represent ~20% of the most abundant genera in Trifolium roots, which was one-quarter of the ~3500 detected OTUs in a manageable effort to increase the cultivated endophytic bacteria from a legume (Table 1).
On the other side, culture-independent methods mostly rely on the extraction of bacterial genetic material from plant tissues. The genomic DNA can then be analyzed using a range of molecular fingerprinting techniques such as Amplified rDNA Restriction Analysis, Gradient Gel Electrophoresis, and Terminal Restriction Fragment Length Polymorphism (RFLP) [16]. In recent years, DNA fingerprinting techniques have been set aside by more advanced molecular techniques. Those new methods involve DNA extraction from the entire bacterial population to sequence a specific phylogenetic marker, such as the 16S rRNA gene, or the whole genome [76]. In addition, using RNA instead of DNA, it is possible to detect active functional diversity, which provides information about the transcriptionally active functions, as well as the massive analysis of proteins (peptides) or metabolites (by high throughput analysis of “omics”). The latter two do not provide taxonomic information but a functional one.
The sequence-based methods allow a deeper analysis of the endophytic diversity than traditional fingerprinting, although some of the species with low abundance might be still missed. To minimize those losses, it is important to sequence with high depth and carry out rarefaction analysis (to check that the OTU versus the diversity or richness reaches the plateau). Other technical considerations for sequencing analysis are discussed in detail by Lucaciu et al. [77].
The bacterial diversity of the microbiome can be described taxonomically and functionally by different approaches. The most traditional strategy is the taxonomic description of the diversity, which identifies the species present in the microbiome and quantifies their abundance by genome or specific gene sequencing. From that data, researchers have started to uncover what is known as the “core microbiome” [78], which is defined as the group of species present in one plant across different genotypes, environments, developmental stages, etc. Depending on the scale of the analysis, a higher or lower number of species are shared among them. For instance, if dicot and monocot species are compared, the number of shared species will be lower than if two cultivars of the same species are compared in the same environment. A core endophytic microbiome of roots of red clover (Trifolium pratense) includes 70% of Rhizobia, and it was dominated by the genera Pantoea, Sphingomonas, Novosphingobium, and Pelomonas [52] (Table 1). Glycine spp. nodules showed a majority of Ensifer genera, followed by Enterobacter, Stenotrophomonas, and Chryseobacterium (>0.5%), and some nonrhizobial bacteria only in soybean (Glycine max), including Enterobacter cloacae (3.62%), Stenotrophomonas sp. CanR-75 (2.79%), and Stenotrophomonas maltophilia (2.41%) [40] (Table 1). Overall, little is known about the core endophytic microbiome in legumes, although some core rhizospheric microbiomes have been described (e.g., [79]).
In addition to the core microbiome, the “keystone” species have been described [80]. Keystones are highly connected species that largely change the structure and function of the microbiome when removed. They may be predicted by co-occurrence networks (by correlation analysis) and are defined as those whose abundance highly correlates with most of the other species [81]. Those correlations can be positive or negative (i.e., two species are always together or the presence of one excludes the other), and the interaction between each other may be indirect (for instance, mediated by a change in the host) [82]. It has been predicted that when the keystone species is missing, the abundance and proportion of the community change, and occasionally, one species may extremely proliferate over the others. Knowing which are the keystone species for one host is critical to effectively design any agricultural management strategy to protect a healthy microbial community and improve the fitness of the crop.
A second strategy to characterize the microbiome is the functional description, based on the metabolic functions present in the microorganisms. According to the previous model (with a core microbiome and keystone species), the communities in the microbiome are built to occupy functional niches [81]. This means that one species might be (at least partially) replaced by another one, which provides the same function to the community and/or the host. Those key functions of a particular species are given by a set of genes that allow the microbe to effectively interact and benefit the rest of the microbial community and the plant host under specific conditions. These functional traits can be screened and studied by any “omic” analysis and then grouped by the presence of specific metabolic functions (see [83, 85] in Table 1). For instance, the most important genes differentially detected in the rhizosphere of pea (Pisum sativum) under different tillage and fertilization treatments were genes coding ABC transporters and secretion systems, transcription factors, peptidases, methane metabolism, quorum sensing, and bacterial motility proteins [85]. To understand which services the microbial community provides and may favor the host plant, the functional analysis may be more useful than a taxonomic-only approach. However, both are necessary and provide valuable information about the microbiomes.
3. Benefits of endophytic microbiota to the host plant
Once within the plant, endophytes might provide several benefits. We grouped them into three different kinds: direct growth promotion, protection against pathogens, and protection against abiotic stress (Figure 1).
Direct promotion occurs when endophytes stimulate shoot and/or root growth by increasing the availability of limiting nutrients or producing compounds that directly stimulate growth. On the other hand, indirect promotion occurs when the endophytes can protect the plant against diseases, pests, or environmental stress, indirectly improving the host performance [86]. The molecular mechanisms and pathways are not exclusive for each direct or indirect growth promotion effect. A single endophytic bacterial strain may have more than one of these plant-growth-promoting traits (e.g. [37, 41, 48, [49, 55] in Table 1, and [56, 57, 63, 65, 66, 68] in Table 2).
3.1 Increase of nutrient availability
The main mineral nutrients required for plant growth are nitrogen, phosphorus, and iron. There are numerous plant-growth-promoting microorganisms able to increase their availability, and some mechanisms have been determined.
3.1.1 Biological nitrogen fixation (BNF)
Nitrogen is crucial for plant growth and health. Approximately 30–50% of the N in crop fields results from BNF by soil microorganisms. The ability to fix atmospheric nitrogen (N2) is present in various bacterial species that are either free-living or endophytically associated with plant roots. BNF is the most and long-term studied plant-growth-promoting effect of soil microorganisms in legumes [87, 88]. Other plant growth promoter bacteria genera, different from rhizobia, are also able to enhance the acquisition of N by legumes. Anzuay et al. [89] and Taurian et al. [90] observed that endophytic bacteria belonging to Serratia, Acinetobacter, Bacillus, and Enterococcus enhanced peanut (Arachis hypogaea) N content. Dey et al. [91] reported that the increase in the number of nodules in plants inoculated with growth promoter bacteria could be attributed to the enhancement of root growth and root length. This enhancement provides more sites for nodulation by rhizobial strains in the soil. Furthermore, since BFN is a highly demanding ATP process, phosphorus is a critical nutrient for legumes.
3.1.2 Phosphate solubilization and mineralization
Even in phosphorus-rich soils (such as phosphate-fertilized soils), most of this element is in insoluble forms, and only a small proportion (~0.1%) is available to plants [92]. The solubilization of phosphates in the rhizosphere is one of the most common modes of action of growth promoter microbes that enhance nutrient availability to plants [93]. Phosphate-mineralizing and phosphate-solubilizing bacteria (PMB/PSB) secrete phosphatases and organic acids to convert insoluble phosphates (organic and inorganic) into soluble monobasic and dibasic ions [93]. Among legume endophytes, there are several phosphate-solubilizing bacteria able to promote plant growth, and some studies demonstrated that plant growth promotion was directly correlated with the increase of P in the plant tissues [89]. Soybean and peanut endophytes solubilize mineral phosphate [90]. In addition, several studies described endophytic bacteria with phosphate-solubilizing/-mineralizing ability that increase legume growth [89, 90, 94, 95]. The inoculation of pea with phosphate-solubilizing Pseudomonas spp. isolated from this legume, enhanced the plant biomass [96]. Pantoea spp. isolated from root nodules of peanut showed a strong phosphate-solubilizing activity [97]. However, the inoculation of phosphate-solubilizing bacteria isolated from peanuts did not promote growth when they were inoculated in the rapeseed culture [98]. These results point to a specific plant-bacteria interaction that directly affects the ability to promote growth or the efficiency of the mechanism.
The main phosphate-solubilizing mechanism in Gram-negative bacteria involves the bacterial PQQ cofactor, described as essential in P nutrition and plant growth. Mutation in the pqqH gene from Pseudomonas fluorescens caused the loss of the phosphate-solubilizing phenotype and plant growth promotion ability on tomato plants [99]. In legumes, Ahmed and Shahab [100] observed that a non-producing-PQQ bacteria (which lost the phosphate solubilization ability) showed a decrease in the growth promotion of bean plants. On the contrary, Ludueña et al. [101] determined that in the non-producing PQQ strain Serratia sp. promoted the growth of peanut at a similar level to the wild type, indicating that PQQ is not essential for growth promotion.
3.1.3 Iron uptake
Iron is essential for all living organisms, and its bioavailability in the soil is limited. Siderophores are small molecular compounds, secreted by microbes, which chelate iron in the soil and generate soluble complexes that can be absorbed by plants [97]. Microbial siderophores’ secretion directly stimulates plant growth by increasing the availability of iron in the soil surrounding the roots [102]. Plants lacking soil bacteria suffered from iron deficiency [103]. Therefore, this mechanism helps plants to thrive in low-iron soils. The inoculation of black mung bean (Vigna radiata) with the siderophore-producing endophyte, Pseudomonas sp. GRP3, reduced iron deficiency and chlorotic symptoms and increased the content of chlorophyll a and b [104]. Furthermore, since diazotrophic organisms require Fe+2 and Mo+2 factors for the functioning and synthesis of nitrogenase, iron solubilization by microbes also improved nitrogen fixation in legumes [105]. Native peanut isolates produce siderophores together with other plant-growth-promoting traits, increasing peanut growth and performance [106].
3.2 Phytostimulators
Endophytic bacteria directly promote plant growth by the production of phytohormones, such as auxin or cytokinin, or by lowering the plant ethylene (ET) levels. By these mechanisms, bacterial endophytes can also accelerate seedling emergence and promote plant establishment under adverse conditions.
3.2.1 Phytohormone-like molecule production
The production of phytohormones-like compounds is considered an important trait of endophytes that positively affects the growth and development of many plants including legumes [8, 10, 107]. Thus, changes in plant growth frequently reflect alterations in phytohormone levels induced by endophytes [3]. But, even when production of these compounds by growth promoter microbes has been demonstrated, that effect cannot be unequivocally attributed to them.
The five main phytohormones produced by bacteria are auxins, cytokinin, gibberellins, ET, and abscisic acid (ABA). It has been postulated that genes encoding biosynthesis of the auxins, cytokinin, and gibberellins are often present in the metagenome of plant endophytic bacterial communities [108]; however, it has not been yet explored in legumes using any omics approach (ET and ABA are discussed in Section 3.4.3).
Among these growth regulators, auxins are the most studied. These compounds affect plant growth by inducing cell enlargement and division, root development, apical dominance, increase growth rate, photo- and geo-tropism [109]. The production of auxin-like compounds increases seed production and germination along with increased shoot growth and tillering. Within these compounds, indole-acetic acid (IAA) is the most frequent and indeed most studied phytohormone in growth promoter bacteria. IAA produced by endophytic bacteria is one of the most relevant and studied effector molecules in growth promotion, pathogen defense, and plant-microbe interactions [104]. For instance, rhizobia from soybean, pea, and faba bean nodules not only fix nitrogen and produce siderophores, but also auxins (see Refs. [54, 110] in Tables 1 and 2, and [61]). IAA can be synthesized directly by plant-associated microbes, and ~ 80% of the rhizosphere bacteria may produce IAA [69, 111]. For instance, it could be produced by Alcaligenes, Azospirillum, Pseudomonas, Pantoea, Rhizobium, and Enterobacter in the presence of L-tryptophan as a precursor, although there are other pathways and a variety of auxins, such as indole-3-butyric acid (IBA), indole-3-pyruvic acid (IPA), or tryptophol (TOL), which are also produced by growth promoter bacteria [112].
Cytokinins are another group of growth-stimulating phytohormones that are responsible for cell division, plant senescence, seed germination, flower and fruit development, and apical dormancy [113, 114]. Although cytokinins are produced by several growth promoter microbes, few studies have demonstrated their beneficial effects.
Gibberellins are involved in many developmental processes in plants, such as flowering regulation, seed germination, stem and leaf elongation [114], but also the promotion of nodule organogenesis and the negative regulation of the rhizobial infection and root system development [115].
Several bacteria produce and regulate the production of more than one phytohormone, such as the rhizobacteria Bacillus aryabhattai, which produces ABA, IAA, cytokinin, and gibberellic acids in vitro and promotes soybean growth [116]. Thus, inoculation with endophytic bacteria may benefit legumes via the production or suppression of some phytohormones.
3.2.2 Volatile compounds and other phytostimulators
Some growth promoters’ bacteria can regulate plant growth by releasing volatile compounds [86]. For instance, B. subtilis, Bacillus amyloliquefaciens, and E. cloacae promote plant growth in legumes by releasing volatiles, such as 2,3-butanediol and acetoin [117, 118], while the mutants of B. amyloliquefaciens IN937a and B. subtilis GB03, blocked in their biosynthesis, did not promote Arabidopsis growth [118]. Studies on growth promotion by Chryseobacterium rhizoplane in mung bean indicate that 2,3-butanediol is the molecule causing growth stimulation [119]. Growth promotion mechanisms of volatiles in plants were reviewed by Sharifi and Ryu [120].
Other nonvolatile molecules such as bacterial cell components or secreted compounds have been proposed to be plant growth stimulators. The endophyte Serratia proteamaculans was able to promote soybean growth by the production of a lipo-chitooligosaccharide [121]. And the PQQ peptide, previously mentioned to be associated with P solubilization, has also shown growth promotion [99], antifungal activity, and the ability to induce systemic resistance [86]. The role of PQQ in plant-microbe interaction has been reviewed by Carreño-Lopez et al. [122].
Lastly, endophytes can generate allelopathic effects inhibiting the growth of neighboring plants or protecting the host plant from allelopathic effects from adjacent plants [123]. For example, endophytic bacteria of red clover seem to be responsible for the negative allelopathic effects observed over maize, reducing seedling emergence and height [124]. Additionally, some weeds have negative allelopathic effects on legumes, mediated by their endophytic bacteria, which inhibit nodulation [125].
Overall, there is a body of evidence that suggests that enhancing or regulating phytohormone or other phytostimulators via endophytic microorganisms is a viable strategy to increased crop production in agriculture [108], and because of these attributes, endophytes have gained ground in the area of agricultural sustainability.
3.3 Protection against pathogens
Among the major factors restraining agriculture are crop diseases and pests, while one important driver of plant health is the structure and dynamics of the plant-associated microbial communities [126]. In recent years, a deeper understanding of the endophytic microbiome and its potential has been achieved to become a fundamental tool in phytosanitary management and reduce the damage of plant diseases.
Endophytes can decrease the harmful effects of pathogens by different mechanisms, including direct and indirect mechanisms [104]. Direct inhibition of pathogens is mainly mediated by the synthesis of inhibitory allelochemicals such as antibiotics, hydrogen cyanide, iron-chelating siderophores [127], secretion of lytic enzymes, or quorum quenching (QQ) by degrading pathogens autoinducer signals [128]. Indirect biocontrol mainly includes the induction of the plant systemic resistance that inhibits the proliferation of a broad spectrum of phytopathogens [129].
3.3.1 Antibiosis
Most endophytes have been reported to produce secondary metabolites, and some of them exhibit antibacterial and antifungal properties, which help to inhibit the growth of phytopathogenic microorganisms [44]. Many metabolites with antimicrobial properties synthesized by endophytes have been described so far, such as flavonoids, peptides, quinones, alkaloids, phenols, steroids, terpenoids, and polyketides. Antimicrobial properties of bacterial metabolites were recently reviewed [130]. Hansen et al. [131] studied the microbiome of alfalfa (Medicago sativa) nodules and identified two families of molecules produced by Brevibacillus brevis in planta, such as antibacterial thyrozidines, and a new set of gramicidin-like molecules, britacidins. They conclude that, in addition to nitrogen fixation, it is likely that legume root nodules are also a source of active antimicrobial production.
3.3.2 Lipopeptides
Lipopeptides are low-molecular-weight cyclic peptides attached to a hydrophobic fatty acid. These molecules are classified into three families: surfactin, iturin, and fengycin. Iturins and fengycins show strong antifungal activities while surfactins exhibit strong antibacterial activity. Antimicrobial lipopeptides can form toroidal-like pores on cell membranes leading to membrane permeation and/or disintegration and protect plants directly suppressing the growth of pathogens or inducing systemic resistance [132]. Recently, 263 different lipopeptides were synthesized by 11 microbial genera, with Bacillus being the most abundant [133].
The common bean root microbiome was used to search potential biocontrol agents of Fusarium sp., Macrophomina sp., and Alternaria sp. fungi, causal agents of root rot disease [65]. Biocontrol assays conducted under controlled conditions demonstrated that B. amyloliquefaciens, B. halotolerans, Bacillus velezensis, Agrobacterium fabrum, and Pseudomonas lini displayed the highest protective effect, and lipopeptide biosynthetic genes encoding surfactin, iturin, bacillomycin, and fengycin were present. These bacteria can produce at least one or more lipopeptides that may be involved in biocontrol activity.
3.3.3 Lytic enzymes
During plant colonization, endophytes produce numerous enzymes, which successively aid the hydrolysis of the plant cell wall. There are numerous types of enzymes such as chitinases, cellulases, hemicellulases, and 1,3-glucanases [70, 134]. These enzymes are also capable of degrading fungal (and oomycetal) cell walls hyphae, spores, and sporangia, thus contributing to the protection of the plant. The isolate Pseudomonas spp. EGN 1 was the most promising bioagent for the management of the stem rot (Sclerotium rolfsii) in groundnut, mediated by an important protease and cellulase production [57]. While, Brigido et al. [135] evaluated the diversity and functionality of the endophytic bacterial strains in the roots of native legumes from two different sites in Portugal, finding 15 isolates with a high cellulase production.
3.3.4 Hydrogen cyanide
A few bacterial species are known to produce and excrete hydrogen cyanide, a potent inhibitor of cytochrome c oxidase and several other metalloenzymes [136]. The host plant is unaffected by the bacteria or the hydrogen cyanide produced by it. For this reason, hydrogen-cyanide-producing bacteria have an application as biological control agent. Zaghloul et al. [137] isolated a total of 167 endophytic bacterial from roots, nodules, leaves, and stems of faba bean (Vicia faba), pea, fenugreek (Trigonella foenumgracum), lupine (Lupinus spp.), common bean (Phaseolus vulgaris), and rice (Oryza sativa) at flowering stage. About 82% of the isolates showed positive results of hydrogen cyanide production. In another recent investigation, ~20 endophytic bacteria isolated from roots and nodules of chickpea (Cicer arietinum) and pea showed HCN production [66].
3.3.5 Siderophores
As previously mentioned, siderophores chelate iron in the soil making it more available for plants. Furthermore, by tightly binding the iron, siderophores reduce its bioavailability for plant pathogens and facilitate the death of the phytopathogens [138]. Some of the siderophores are known to be produced by endophytes, such as hydroxymate, phenolate, and/or catecholate types, confer biocontrol activities [139]. Also, the role of siderophores as part of the protective effect of the induced systemic resistance has been described in many studies. The production of siderophores is very common among Pseudomonas, Frankia, Streptomyces sp. Several researchers described endophytic bacteria producing siderophores isolated from different legumes as peanut, faba bean, soybeans, chickpea, pea, and bean [65, 66]. Bahroun et al., [140] demonstrate that Rahnella aquatilis B16C, Pseudomonas yamanorum B12, and P. fluorescens B8P isolated from faba bean nodules suppressed Fusarium solani root rot in three faba bean cultivars in greenhouse. The three strains were able to produce siderophores and significantly reduced the disease severity. Zhao et al. [54] obtained 276 isolates from root nodules of soybean, six of which showed antagonistic to the pathogenic fungus Phytophthora sojae 01. The isolates were identified as Enterobacter, Acinetobacter, Pseudomonas, Ochrobactrum, and Bacillus genera. The high correlation of siderophores production and the fungal inhibition of nodule endophytic bacteria in that study supported the idea that the ferrous absorption by endophytic bacteria may be a viable inhibitory mechanism.
3.3.6 Quorum quenching
The regulation of gene expression in response to fluctuations in cell-population density is known as “quorum sensing.” Many important bacterial processes are regulated by it. Quorum sensing regulates gene expression depending on the accumulation of a signal molecule in the environment. The signal, called autoinducer, allows the bacteria to perceive the existing population density and jointly executed responses. Gram-negative bacteria use acyl-homoserine lactone (AHL) as an autoinducer, whereas Gram-positive bacteria utilize modified peptides [141]. The bacterial quorum sensing controls a wide variety of physiological processes such as virulence, extracellular polymeric substances (EPS) production, mobility, and biofilm formation among others, which are essential for the establishment of a pathogen in the host plant [142].
Often endophytic bacteria can disrupt quorum sensing. This ability to interfere with bacterial cell-to-cell communication was collectively called “quorum quenching” and can be crucial to prevent the plant colonization by pathogenic bacteria that use quorum sensing to coordinate virulence [143]. Several chemicals and enzymes have been identified that target the key components of bacterial quorum-sensing systems in the recent years (such as [33]). The mechanisms of quorum quenching may be the inhibition of the signal synthesis or detection, signal enzymatic degradation (by enzymes such as AHL acylase, AHL lactonase, and oxidoreductases), or synthesis of structural analogs of the signal [144]. Lopes et al. [145] reported antimicrobial activity against Pseudomonas syringae pv. tabaci or Hafnia alvei 071 in endophytic bacteria isolated from common bean. The isolates Microbacterium testaceum BAC1065, BAC1100, and BAC2153, Bacillus thuringiensis BAC3151, and Rhodococcus erythropolis BAC2162 exhibited a greater ability to inhibit the response of AHL reporter.
3.3.7 Insecticides
Some metabolites with insecticidal action have been described. The famous B. thuringiensis produces crystalline inclusion bodies consisting of delta-endotoxins (also referred to as Cry proteins) during sporulation. These proteins, which are formed by variable-molecular-weight polypeptides (27–140 kDa), are highly toxic for a broad range of pest insects [146]. P. fluorescens strains exhibited a protective effect against aphids and some herbivorous beetles and termites [147]. The bacterium Lysinibacillus sphaericus (former Bacillus sphaericus) produces sphaericolysin, which is toxic for Spodoptera litura [148].
3.3.8 Induction of systemic response
Induced systemic resistance (ISR) is a term used for the resistance stimulated by chemicals agents or signals (elicitors) produced by beneficial microorganisms [149], whereby the plant’s innate defenses are potentiated against subsequent biotic challenges. In this way, the endophytes enhance the plant defenses against many pathogens [129]. The plant hormones jasmonic acid (JA) and ET are responsible for the regulation of the group of interrelated signaling pathways required to activate ISR. The main routes by which microbes regulate ISR in plants include: (i) phytohormones, (ii) pathogen-associated molecular patterns (PAMPs)/microbe-associated molecular patterns (MAMPs), and (iii) several elicitors (volatile organic compounds, siderophores, phytases, miRNAs, among others) [150]. Bacterial endophyte-mediated ISR has a broad spectrum of effectiveness. It was demonstrated that Acinetobacter, Azospirillum, Rhizobium, Pseudomonas, and Bacillus are beneficial inducers of systemic resistance in both leguminous and nonleguminous plants [151]. Dey et al. [91] described an endophytic isolate Klebsiella pneumoniae HR1 from the root nodules of black mung bean (Vigna mungo) capable of reducing the occurrence of Macrophomina phaseolina, which is the causal agent of the root rot disease in Vigna. The lowest percentage of disease incidence (18.2%) was observed when K. pneumoniae was applied in dual mode (seed bacterization + soil drench application). The increased activities of peroxidase (PR9), chitinase (PR3), and β-1,3-glucanase (PR2) in leaves indicated that K. pneumoniae HR1 induces a systemic response.
Endophytic bacteria have diverse mechanisms that could contribute, even simultaneously, to protect the plant against the attack of different pathogens, having the potential to produce a more efficient pathogen control on the fields.
3.4 Abiotic stress tolerance
Under abiotic stress conditions (such as drought, salinity, flooding, heat, chilling, or heavy metals), several metabolic responses are shared among plant species. Most of the stresses cause photosynthesis inhibition, oxidative stress, and hormone imbalances ending in reductions of shoot growth and yield impairments [10, 97, 152, 153, 154]. In addition, some of the responses are interconnected, for instance, reactive oxygen species and hormones mutually affect each other at early and late phases of abiotic stress (reviewed by [155]).
Endophytic bacteria can protect the host plant against some of those deleterious effects, by at least two different ways (alone or combined): (i) activation of host stress response systems soon after exposure to stress (named induced systemic tolerance), and (ii) biosynthesis of chemicals, which will contribute to the stress tolerance in the host [9]. Here we focus on three mechanisms by which the bacteria can protect the plant host against abiotic stress: redox status, water balance, and hormone regulation.
3.4.1 Redox status regulation
Oxidative damage (caused by reactive oxygen and nitrogen species) is a common consequence of environmental stress, which may cause damage to lipids, proteins, and overall to any subcellular component [156]. Then, the activation of the enzymatic and nonenzymatic antioxidant system is critical to tolerate adverse conditions. Several endophytic bacteria mediate a higher induction of the antioxidant system under stress. For instance, under salinity, the inoculation of peanut with the halotolerant bacteria Brachybacterium saurashtrense JG-06, Brevibacterium casei JG-08, or Haererohalobacter JG-11 showed lower oxidative damage, ion leakage, and K/Na ratio and higher growth, IAA, and Ca [157], while the inoculation of B. subtilis (alone or combined with Mesorhizobium ciceri) of chickpea reduced hydrogen peroxide accumulation and improved plant growth [10]. Soybean plant inoculated with Curtobacterium sp. SAK1 induced polyphenol oxidase activity, associated with growth protection and hormonal changes [158], while inoculated with Pseudomonas simiae increased catalase and peroxidase, but not polyphenol oxidase gene expression under salinity [159]. Also, soybean inoculated with B. cereus, Pseudomonas otitidis, and Pseudomonas sp. showed a reduction of hydrogen peroxide and membrane oxidative damage caused by PEG-induced drought [160]. However, if these responses are generated by the plant or bacterial enzymes remains unknown.
3.4.2 Water use efficiency regulation
Under stress, plant tissues usually modulate osmotic and water retention, by stomata activity and/or accumulation of osmotically active compounds. The latter compounds, also known as compatible solutes, include sugars (e.g., sucrose, trehalose, etc.), organic acids (e.g., malate), inorganic ions (e.g., calcium), amino acids (e.g., glycine betaine, proline) [161]. An increase in drought tolerance was detected after the inoculation of Sphingomonas sp. LK11 (isolated from Tephrosia apollinea) in soybean, by the accumulation of sugars and amino acids (glycine, glutamate, and proline) [162], and after the inoculation with Rhizobium etli in common bean, by the overexpression of trehalose-6-phosphate synthase [163]. Trehalose is an osmotically active compound that accumulates both in plants and microbes under stress. In particular, the role of trehalose in the tripartite symbiosis between plants, rhizobia, and arbuscular mycorrhiza under abiotic stress has been recently reviewed [164].
The optimal regulation of water use efficiency is critical to improved crop production. On one side is essential to survive dehydration stress (such as drought, salinity, heat, and chilling), but a constitutively highly efficient water use may reduce yields, by reducing CO2 assimilation. The use of bacteria that contribute to transiently intensify stress-tolerance responses can help to improve productivity in marginal environments. In addition, if the endophytic bacteria enhance the osmocompatible compounds in response to the stress, it is possible to increase not only the tolerance to drought, but also the tolerance to chilling, heat, and salinity stress, which share a “dehydration” component. In the latter case, we expect a partial tolerance due to the ion toxicity, not related to the reduction in water potential.
3.4.3 Hormone regulation
As it was mentioned before, endophytic bacteria can regulate hormone synthesis and degradation and synthesize some of the plant hormone-like compounds by themselves. In addition, specific hormone regulation could also protect against abiotic stress increasing growth, yield, and survival.
Abscisic acid (ABA) is the main plant hormone related to water stress. It stimulates root growth and optimizes water uptake and nutrient acquisition, regulates shoot and root hydraulic conductivity, and upregulates the antioxidant system and compatible osmolytes synthesis [161]. The inoculation of Sphingomonas in soybean leaves induced ABA accumulation and reduced chlorophyll degradation and growth inhibition. However, under drought, ABA levels were lower in inoculated plants. So, in this case, the initial increase of ABA might have a role in acclimation to the stress induced by the bacteria inoculation [162]. In addition, ABA may interfere with SA-, JA-, and ET-mediated plant defenses [165], which may have undesired consequences under biotic stress.
Ethylene (ET) is usually considered a plant growth inhibitor, but at low levels, it can promote growth in several plant species. At moderate levels, ET inhibits both root and shoots elongation, while at high levels, enhances senescence and organ abscission [166]. The direct precursor of ET in the plant biosynthetic pathway, 1-aminocyclopropane-1-carboxylate (ACC), is exuded from plant roots together with other amino acids. The enzyme ACC deaminase cleaves ACC into ammonia and alfa-ketobutyrate. Plant growth promoter bacteria that express the enzyme ACC deaminase utilize their products (ammonia and ketobutyrate) as nitrogen and carbon sources, respectively. Bacterial ACC deaminase is not excreted from the bacterial cytoplasm [167]; hence, the decrease of plant ET levels relies on the ability of ACC deaminase expressing bacteria to take up ACC before it is oxidized by the plant’s ACC oxidase [167]. When those bacteria are present, ET production could be lowered, relieving stress-induced growth inhibition [168]. For instance, the inoculation of pea (P. vulgaris) plants with Aneurinibacillus aneurinilyticus and Paenibacillus sp., two strains with high ACC activity in vitro, increased salt and drought tolerance. The combined inoculation reduced plant ET content and increased root and shoot length and biomass, as well as chlorophyll content [169]. The inoculation of alfalfa plants with Bacillus megaterium NMp082, which can produce ACC deaminase activity and IAA in vitro, also enhanced their salt tolerance [170]. Lastly, a novel mechanism was proposed in which salt tolerance is mediated by the activation of ET signaling. The inoculation of alfalfa with the bacteria Enterobacter sp. SA187 (isolated from a desert plant) increases salt tolerance, and studies in Arabidopsis indicate that the bacteria activate the ET signaling pathway [171]. The different mechanisms by which microorganisms can interfere with ET signaling were reviewed by Ravanbakhsh et al. [167].
Auxins regulate many important physiological processes related to growth and development affecting photosynthesis and responses to stress [161]. Under stress, auxins stimulate root elongation and density, increasing the water and nutrient availability, although they may interfere with SA-dependent plant defenses.
The inoculation of chickpea with Serratia sp. in nutrient-deficient soil induced more IAA and higher yields [172], while the same plant inoculated with IAA-producing B. subtilis NUU4 in combination with M. ciceri IC53 stimulated root and shoot biomass and improved nodule formation under salt stress [173]. Soybean plants inoculated with B. aryabhattai strain SRB02, which produces IAA, GA, and ABA, showed higher drought tolerance through stomatal closure, and higher root and shoot rates under high temperatures [116], and the same host treated with Sphingomonas sp. LK11 and Serratia marcescens TP1 (which produced IAA in vitro) stimulated root and shoot growth with increased ABA and GA and reduction of JA [162]. Overall, abiotic stress protection mediated by plant hormones and crop salinity protection mediated by beneficial bacteria have been reviewed [10, 174, 175].
Some primary stresses share the responses among them, such as those that generate dehydration (water or temperature deficit) or oxidative stress (dehydration, hypoxia, ions). For example, the double inoculation of chickpea with M. ciceri IC53 and B. subtilis NUU4 reduced the infection rate of root rot caused by Fusarium solaniin salty soils [173], although the mechanism was not determined. Then, a bacteria strain, inducing a protective mechanism against oxidative stress, can protect the crop against a diversity of stress, which generates redox imbalances. Consequently, knowing the responses that each stress triggers in the plant may allow us to predict which bacteria or group of them could protect the plant against a combination of stresses.
4. Synthetic communities of plant-associated bacteria to a more sustainable agriculture
Natural microbial communities within the plants are complex systems, with unknown functions and interrelationships among the microbial species and with the host plant. Small consortia of bacteria, with a “designed” composition, called “synthetic communities,” reduce the complexity of those systems to be studied and used. The goal is to simplify the network while preserving the interactions and most of the functions, which may be lost in single plant-microbe interactions [175]. The use of synthetic communities allow us to ask questions about the performance and stability of the microbial community as well as to study conditions necessary to generate interaction patterns required to provide specific benefits. They are not only valuable as models but also as assays for biotechnological approaches [176].
4.1 How to study synthetic communities?
Manipulative experiments with synthetic bacterial communities can validate the predicted keystone species and, in general, help to find out specific effects of the resulting community under some pathogen infection or environmental condition. Those studies required in vitro experiments in gnotobiotic (germ-free) systems [11], where the plant is inoculated with a few or several microbial species, and the diversity is monitored across time. For instance, a gnotobiotic system was used to study the bacteria-colonizing alfalfa nodules [131]. The authors inoculate alfalfa with the four accessory bacterial members B. brevis Ag35, Paenibacillus sp. Ag47, Pseudomonas sp. Ag54, and Pantoea agglomerans Ag15, plus the nodulating strain Sinorhizobium meliloti RM1021. They observed that the addition of B. brevis neutralized the cooperation between Pseudomonas sp. Ag54 and Paenibacillus sp. Ag47, shifting the community from cooperative to competitive.
Another alternative, it is to use synthetic communities in a non-germ-free environment (more accessible and simpler to set up) to evaluate the protective or antagonist effect of a small group of species under a particular condition. Overall, only a few studies of the kind have been carried out in legumes until now. For instance, Lu et al. [177] described the diversity of nonrhizobial bacteria (32 genera) in legume nodules inoculated with Bradyrhizobium elkanii H255, Rhizobium multihospitium–like HT221, or Burkholderia pyrrocinia with or without the addition of N fertilization. The study suggested a vital role of that group of bacteria in N fixation in legumes.
The synthetic communities are a way to understand how microbial communities are built in the plants but also the base to a more complex (and likely more effective) phytostimulation effects, biological control of diseases, and protection against abiotic stress.
4.2 Can we manipulate the plant microbiome to improve the fitness or yield of legumes?
There are a variety of strategies to manipulate the microbiome of a plant host and could be classified according to the direct target: (i) the microbiome itself, (ii) the plant genome, or (iii) the holobiome (plant plus microbial community) (reviewed by [39, 178]).
The microbiome (i) can be modified by the exogenous inoculation of the microbe, increasing the abundance of a single strain or a few species together. The first case is the most traditionally used, and there are thousands of examples, such as the inoculation with rhizobia. In those cases, the single strain should be compatible with the host genotype and able to overcome the competence of the native microbiome and the environmental conditions. The second case is open to unexplored scenarios, such as an infinite possibility of a higher number of strains/species combinations. This strategy is just starting to be explored, such as with non-nodulating bacterial species present in the nodules (and sometimes in the rest of the plant) that promote nodulation. For instance, the inoculation of common bean (P. vulgaris L.) with Paenibacillus polymyxa and B. megaterium strains showed a synergistic effect with Rhizobium strains on the plant growth [179]. On the contrary, the inoculation of alfalfa with different strains of the mutualistic P. fluorescens, showed that the increase in the community richness led to a negative complementary effect causing the loss of the protective effect against pathogens [180]. These results highlight the importance to evaluate the effects of any agricultural treatment or management on the microbial community.
The inoculation with synthetic communities has the advantage (over the use of the native microbiome) to allow the design of a community, which includes distant species (which may provide complementary benefits), or similar species, which increase the efficiency of the community (by using a wider diversity of resources) [19]. However, with the number and diversity of species, it also increases the complexity to handle the system and to commercialize the inoculants.
The plant genome (ii) could be manipulated by traditional breeding, gene editing, or transgenesis, changing the ability of the host to interact with the microbes (such as changing the exudates or volatiles). Instead of only breeding for pathogen resistance or abiotic stress tolerance, this could be a complementary alternative to select crop legumes to be more responsive to the presence of beneficial microbes [181]. For instance, modern accessions of common bean showed a lower abundance of Bacteroidetes and higher of Actinobacteria and Proteobacteria than the wild accession [79], with a gain in the diversity of rhizospheric bacterial and a stronger effect of the bean genotype [182]. In addition, Mendes et al. [183] showed that common bean breeding for Fusarium oxysporum resistance altered the functionality of the rhizosphere, unintentionally increasing the host protection against other pathogens. We hypothesize that a similar effect is happening in the endosphere, although it has not been explored yet. Additionally, when using this approach, it is relevant to evaluate that host defenses against pathogens are still functional.
Lastly, the holobiome (iii) could be altered through specific agricultural practices such as crop rotation, mineral, and organic fertilization, tillage practices, etc., favoring a specific community composition or function. Several studies reported the effect of agricultural management on the rhizosphere of legumes and its effect on crop performance. A meta-study showed the effect of crop rotation, intercropping, or companion planting on the rhizospheric microbial richness and diversity [184]. Those agricultural practices did not always have positive effects in richness and diversity, and legume-cereal crop rotation (relevant to reduce N fertilization) showed inconsistent results on the microbiome. A recent study showed that pea-wheat rotations showed no effect in the diversity index, but they affected the specific co-occurrence networks for each crop [185] suggesting a more complex effect of crop rotation that needs to be further studied. Certain chickpea cultivars select a more beneficial microbiome for the subsequent wheat plants, and they were associated with the antagonist species Penicillium canescens [186]. Red clover and potato crops in rotation shared 73% of the bacterial endophytes, and 21% of those species promoted plant growth and yield in potato bioassays [187], while 74% of the shared species showed some degree of in vitro antibiosis against Rhizoctonia solani, a pathogen of both crops. We hypothesize that changing the rhizosphere will affect the endosphere too, by changing the available microbial pool, but that effect has not been explored at legume endophytic microbiomes.
4.3 Are there collateral impacts of using synthetic communities in agriculture?
Lastly, it is important to consider alive microbes will be released to the environment and into products used or consumed by humans and animals, so the potential risks need to be considered and tested [188]. There is no internationally agreed protocol to be complimented, but recently, Vilchez et al. [189] have proposed an Environmental and Human Safety Index (EHSI) protocol to determine the safety of the bacterial strains. The protocol evaluates microbial and animal sensitivity/pathogenicity and ecotoxicity in different model organisms, and it has been validated for many well-known bacteria. In addition, on the agronomical level, little information is available on the nontarget effects on microbial communities and the resulting impact on the soil function [32].
5. Final remarks and future directions
Agricultural legume crops are usually treated with synthetic chemicals to increase growth, control diseases, and mitigate environmental stress, which has high economic, environmental, and health costs. However, there is a myriad of endophytic bacteria that colonize the plant at least in part of its life cycle that could replace or complement those chemicals with great benefits for the plants. In addition, the huge bacterial diversity could be combined to provide several benefits at the same time. For that purpose, the use of synthetic communities is critical to study how the microbial community evolves within the plant as much as their beneficial effects.
The use of synthetic bacterial communities to improve and make more sustainable legume production is still in early stages of development, but it is a promising field. Using synthetic communities has the theoretical advantage of combining strain benefits and contributing to the survival of the bacteria on the field and inside the plant while producing a package of benefits for the legume. Although it is expected to have more difficulties at the time of commercial production.
On the other hand, changes in the agricultural management with some specific purpose could be a more affordable strategy for most of the small-scale producers in low-income countries, which are the ones in more need of sustainable and accessible technologies. Additionally, the use of soil-native microorganisms could have the advantage to reduce possible adverse consequences on the environment and health.
For the moment, the knowledge about endophytic bacteria in legumes, the possibility to “design” synthetic communities for a specific goal, and to manipulate the holobiome by agricultural practices is still incipient. However, the potential benefits for current agriculture to improve yields and sustainability have a great unexplored potential in the endophytic bacterial microbiome of legume crops.
Acknowledgments
This work was supported by FONCyT (PICT STARTUP 2018-0065, PICT 2018-01326) and INTA (I069, I127, I516). CC is a CONICET fellow. MIM and TT are Career Investigators of CONICET. MIM, LV, and MCG are Career Investigators of INTA.
Conflict of interest
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Author contributions
LV and MM conceived and planned the overall idea of the review manuscript. All authors contributed to the article and approved the submitted version.
\n',keywords:"sustainable agriculture, abiotic and biotic stresses protection, food security, endophytic bacteria, synthetic communities",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/80641.pdf",chapterXML:"https://mts.intechopen.com/source/xml/80641.xml",downloadPdfUrl:"/chapter/pdf-download/80641",previewPdfUrl:"/chapter/pdf-preview/80641",totalDownloads:53,totalViews:0,totalCrossrefCites:0,dateSubmitted:"September 19th 2021",dateReviewed:"January 7th 2022",datePrePublished:"March 31st 2022",datePublished:null,dateFinished:"February 27th 2022",readingETA:"0",abstract:"Plant-associated microbiomes confer fitness advantages to the plant host by growth promotion through different mechanisms including nutrient uptake, phytohormones production, resistance to pathogens, and stress tolerance. These effects of the potentially beneficial microbes have been used in a diversity of biotechnological approaches to improve crop performance applying individual bacterial cultures. However, healthy plants host a diversity of microorganisms (microbiota). Next-generation sequencing technologies have offered insights into the relative abundances of different phylogenetic groups in a community and the metabolic and physiological potential of its members. In the last decade, researchers have started to explore the possibilities to use temporal and functional combinations of those bacteria in the form of synthetic communities. In this chapter, we review the benefits of using endophytic bacteria in legumes, the available methodological approaches to study the effects of bacterial communities, and the most recent findings using synthetic communities to improve the performance of legume crops.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/80641",risUrl:"/chapter/ris/80641",signatures:"Mariela I. Monteoliva, Lucio Valetti, Tania Taurian, Clara S. Crociara and María Carla Guzzo",book:{id:"10749",type:"book",title:"Legumes Research - Volume 1",subtitle:null,fullTitle:"Legumes Research - Volume 1",slug:null,publishedDate:null,bookSignature:"Dr. Jose Carlos Jimenez-Lopez and Dr. Alfonso Clemente",coverURL:"https://cdn.intechopen.com/books/images_new/10749.jpg",licenceType:"CC BY 3.0",editedByType:null,isbn:"978-1-83969-491-2",printIsbn:"978-1-83969-490-5",pdfIsbn:"978-1-83969-492-9",isAvailableForWebshopOrdering:!0,editors:[{id:"33993",title:"Dr.",name:"Jose Carlos",middleName:null,surname:"Jimenez-Lopez",slug:"jose-carlos-jimenez-lopez",fullName:"Jose Carlos Jimenez-Lopez"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}},authors:null,sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Plant endophytic microbiome",level:"1"},{id:"sec_2_2",title:"2.1 Colonization and distribution within the host plant",level:"2"},{id:"sec_3_2",title:"2.2 Endophytic bacterial diversity",level:"2"},{id:"sec_4_2",title:"2.3 Factors affecting diversity",level:"2"},{id:"sec_5_2",title:"2.4 How to study microbiome diversity",level:"2"},{id:"sec_7",title:"3. Benefits of endophytic microbiota to the host plant",level:"1"},{id:"sec_7_2",title:"3.1 Increase of nutrient availability",level:"2"},{id:"sec_7_3",title:"3.1.1 Biological nitrogen fixation (BNF)",level:"3"},{id:"sec_8_3",title:"3.1.2 Phosphate solubilization and mineralization",level:"3"},{id:"sec_9_3",title:"3.1.3 Iron uptake",level:"3"},{id:"sec_11_2",title:"3.2 Phytostimulators",level:"2"},{id:"sec_11_3",title:"3.2.1 Phytohormone-like molecule production",level:"3"},{id:"sec_12_3",title:"3.2.2 Volatile compounds and other phytostimulators",level:"3"},{id:"sec_14_2",title:"3.3 Protection against pathogens",level:"2"},{id:"sec_14_3",title:"3.3.1 Antibiosis",level:"3"},{id:"sec_15_3",title:"3.3.2 Lipopeptides",level:"3"},{id:"sec_16_3",title:"3.3.3 Lytic enzymes",level:"3"},{id:"sec_17_3",title:"3.3.4 Hydrogen cyanide",level:"3"},{id:"sec_18_3",title:"3.3.5 Siderophores",level:"3"},{id:"sec_19_3",title:"3.3.6 Quorum quenching",level:"3"},{id:"sec_20_3",title:"3.3.7 Insecticides",level:"3"},{id:"sec_21_3",title:"3.3.8 Induction of systemic response",level:"3"},{id:"sec_23_2",title:"3.4 Abiotic stress tolerance",level:"2"},{id:"sec_23_3",title:"3.4.1 Redox status regulation",level:"3"},{id:"sec_24_3",title:"3.4.2 Water use efficiency regulation",level:"3"},{id:"sec_25_3",title:"3.4.3 Hormone regulation",level:"3"},{id:"sec_28",title:"4. Synthetic communities of plant-associated bacteria to a more sustainable agriculture",level:"1"},{id:"sec_28_2",title:"4.1 How to study synthetic communities?",level:"2"},{id:"sec_29_2",title:"4.2 Can we manipulate the plant microbiome to improve the fitness or yield of legumes?",level:"2"},{id:"sec_30_2",title:"4.3 Are there collateral impacts of using synthetic communities in agriculture?",level:"2"},{id:"sec_32",title:"5. Final remarks and future directions",level:"1"},{id:"sec_33",title:"Acknowledgments",level:"1"},{id:"sec_37",title:"Conflict of interest",level:"1"},{id:"sec_33",title:"Author contributions",level:"1"}],chapterReferences:[{id:"B1",body:'Hallmann J, Quadt-Hallmann A, Mahaffee WF, Kloepper JW. Bacterial endophytes in agricultural crops. Canadian Journal of Microbiology. 1997;43:895-914'},{id:"B2",body:'Santoyo G, Moreno-Hagelsieb G, del Carmen Orozco-Mosqueda M, Glick BR. Plant growth-promoting bacterial endophytes. 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Chickpea genotypes shape the soil microbiome and affect the establishment of the subsequent durum wheat crop in the semiarid North American Great Plains. Soil Biology and Biochemistry. 2013;63:129-141'},{id:"B188",body:'Sturz AV, Christie BR, Matheson BG. Associations of bacterial endophyte populations from red clover and potato crops with potential for beneficial allelopathy. Canadian Journal of Microbiology. 1998;44:162-167'},{id:"B189",body:'Zhang J, Cook J, Nearing JT, Zhang J, Raudonis R, Glick BR, et al. Harnessing the plant microbiome to promote the growth of agricultural crops. Microbiological Research. 2021;245:126690'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Mariela I. Monteoliva",address:"monteoliva.mariela@inta.gob.ar",affiliation:'
Plant Physiology and Genetic Resources Institute—Agricultural Studies Unit (IFRGV—UDEA), National Institute of Agricultural Technology—National Council of Scientific and Technical Research (INTA—CONICET), Argentina
'}],corrections:null},book:{id:"10749",type:"book",title:"Legumes Research - Volume 1",subtitle:null,fullTitle:"Legumes Research - Volume 1",slug:null,publishedDate:null,bookSignature:"Dr. Jose Carlos Jimenez-Lopez and Dr. Alfonso Clemente",coverURL:"https://cdn.intechopen.com/books/images_new/10749.jpg",licenceType:"CC BY 3.0",editedByType:null,isbn:"978-1-83969-491-2",printIsbn:"978-1-83969-490-5",pdfIsbn:"978-1-83969-492-9",isAvailableForWebshopOrdering:!0,editors:[{id:"33993",title:"Dr.",name:"Jose Carlos",middleName:null,surname:"Jimenez-Lopez",slug:"jose-carlos-jimenez-lopez",fullName:"Jose Carlos Jimenez-Lopez"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}}},profile:{item:{id:"300200",title:"Dr.",name:"Petra",middleName:null,surname:"Barisic",email:"pbarisic@efzg.hr",fullName:"Petra Barisic",slug:"petra-barisic",position:null,biography:null,institutionString:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",totalCites:0,totalChapterViews:"0",outsideEditionCount:0,totalAuthoredChapters:"1",totalEditedBooks:"0",personalWebsiteURL:null,twitterURL:null,linkedinURL:null,institution:null},booksEdited:[],chaptersAuthored:[{id:"67963",title:"Nonfinancial Reporting: Theoretical and Empirical Evidence",slug:"nonfinancial-reporting-theoretical-and-empirical-evidence",abstract:"Nonfinancial reporting (NFR) is a relatively new topic in the business practice; it evolved a couple of decades ago. Initially, NFR was mostly disclosed on a voluntary basis. Because of deeper awareness regarding climate change and environmental challenges, as well as pressure from investors, customers, and competition, nonfinancial reporting developed from a voluntary to a mandatory highly standardized practice. This research sought to address the following questions: How understanding of business value creation determines business reporting? How sustainability approach manifests on the company level? What is nonfinancial reporting and why should companies care about it? What are the motivations and benefits for companies and for whom do they publish sustainability reports? What about experiences in public sector? How contemporary concepts of green, circular, and zero-waste economy influence business reporting? Which open questions do organizations face on the path of sustainability reporting? This study contributes to the discussion on NFR and stimulates paradigm shift from profitability toward sustainability as adopting a holistic perspective, respecting people and the planet. 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10. IntechOpen Advertising Sales department makes the decisions about the types of advertisements to include or exclude. Placement of advertising is at the discretion of IntechOpen. IntechOpen retains the right to reject and/or request modifications to the advertisement. An advertisement that is visible online, will be withdrawn from the site at any time if the Editor(s) or Author(s) request its removal.
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Exoplanet characteristics and their comparison to Solar System planets are provided as well as general detection methods and planned probes to gather additional data.",book:{id:"10210",slug:"solar-system-planets-and-exoplanets",title:"Solar System Planets and Exoplanets",fullTitle:"Solar System Planets and Exoplanets"},signatures:"Joseph Bevelacqua",authors:[{id:"115462",title:"Dr.",name:"Joseph",middleName:"John",surname:"Bevelacqua",slug:"joseph-bevelacqua",fullName:"Joseph Bevelacqua"}]},{id:"65725",title:"On the Deviation of the Lunar Center of Mass to the East: Two Possible Mechanisms Based on Evolution of the Orbit and Rounding Off the Shape of the Moon",slug:"on-the-deviation-of-the-lunar-center-of-mass-to-the-east-two-possible-mechanisms-based-on-evolution-",totalDownloads:984,totalCrossrefCites:0,totalDimensionsCites:0,abstract:"It is known that the Moon’s center of mass (COM) does not coincide with the geometric center of figure (COF) and the line “COF/COM” is not directed to the center of the Earth, but deviates from it to the South-East. Here, we discuss two mechanisms to explain the deviation of the lunar COM to the East from the mean direction to Earth. The first mechanism considers the secular evolution of the Moon’s orbit, using the effect of the preferred orientation of the satellite with synchronous rotation to the second (empty) orbital focus. It is established that only the scenario with an increase in the orbital eccentricity e leads to the required displacement of the lunar COM to the East. It is important that high-precision calculations confirm an increase e in our era. In order to fully explain the shift of the lunar COM to the East, a second mechanism was developed that takes into account the influence of tidal changes in the shape of the Moon at its gradual removal from the Earth. The second mechanism predicts that the elongation of the lunar figure in the early era was significant. As a result, it was found that the Moon could have been formed in the annular zone at a distance of 3–4 radii of the modern Earth.",book:{id:"8444",slug:"lunar-science",title:"Lunar Science",fullTitle:"Lunar Science"},signatures:"Boris P. Kondratyev",authors:[{id:"277909",title:"Prof.",name:"Boris",middleName:"Petrovich",surname:"Kondratyev",slug:"boris-kondratyev",fullName:"Boris Kondratyev"}]},{id:"68357",title:"Solar System Exploration Augmented by In Situ Resource Utilization: System Analyses, Vehicles, and Moon Bases for Saturn Exploration",slug:"solar-system-exploration-augmented-by-in-situ-resource-utilization-system-analyses-vehicles-and-moon",totalDownloads:824,totalCrossrefCites:0,totalDimensionsCites:0,abstract:"Human and robotic missions to Saturn are presented and analyzed with a range of propulsion options. Historical studies of space exploration, planetary spacecraft and astronomy, in situ resource utilization (ISRU), and industrialization all point to the vastness of natural resources in the solar system. Advanced propulsion is benefitted from these resources in many ways. While advanced propulsion systems were proposed in these historical studies, further investigation of nuclear options using high-power nuclear electric and nuclear pulse propulsion as well as advanced chemical propulsion can significantly enhance these scenarios. Updated analyses based on these historical visions are presented. At Saturn, nuclear pulse propulsion with alternate propellant feed systems and Saturn moon exploration with chemical propulsion and nuclear electric propulsion options are discussed. Issues with using in situ resource utilization on Saturn’s moons are discussed. At Saturn, the best locations for exploration and the use of the moons as central locations for Saturn moon exploration are assessed. Environmental issues on Titan’s surface may present extreme challenges for some ISRU processes. In-space bases for moon-orbiting propellant processing and ground-based processing will be assessed.",book:{id:"7338",slug:"planetology-future-explorations",title:"Planetology",fullTitle:"Planetology - Future Explorations"},signatures:"Bryan Palaszewski",authors:[{id:"279275",title:"M.Sc.",name:"Bryan",middleName:null,surname:"Palaszewski",slug:"bryan-palaszewski",fullName:"Bryan Palaszewski"}]},{id:"65534",title:"Solar System Exploration Augmented by In Situ Resource Utilization: Lunar Base Issues",slug:"solar-system-exploration-augmented-by-in-situ-resource-utilization-lunar-base-issues",totalDownloads:1108,totalCrossrefCites:1,totalDimensionsCites:1,abstract:"Creating a presence and an industrial capability on the Moon is essential for the development of humankind. There are many historical study results that have identified and quantified the lunar resources and analyzed the methods of obtaining and employing those resources. The idea of finding, obtaining, and using these materials is called in situ resource utilization (ISRU). The ISRU research and development efforts have led to new ideas in rocket propulsion. 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Energetic charged particles impacting materials create electronic excitations, atomic displacements, and nuclear fragmentation. Monte Carlo particle transport simulations are the most common approach for modeling radiation damage in materials. However, radiation damage is a multiscale problem, both in time and in length, an aspect treated by the Monte Carlo simulations only to a limited extent. In this chapter, after introducing the Monte Carlo particle transport method, we present a multiscale approach to study different stages of radiation damage which allows for the synergy between the electronic and nuclear effects induced in materials. We focus on cumulative displacement effects induced by radiation below the regime of hadronic interactions. We then discuss selected studies of radiation damage in materials of importance and potential use for the exploration and settlement on the Moon, ranging from semiconductors to alloys and from polymers to the natural regolith. Additionally, we overview some of the novel materials with outstanding properties, such as low weight, increased radiation resistance, and self-healing capabilities with a potential to reduce mission costs and improve prospects for extended human exploration of extraterrestrial bodies.",book:{id:"10955",title:"Lunar Science - Habitat and Humans",coverURL:"https://cdn.intechopen.com/books/images_new/10955.jpg"},signatures:"Natalia E. Koval, Bin Gu, Daniel Muñoz-Santiburcio and Fabiana Da Pieve"},{id:"80241",title:"The Evolution of the Moon’s Orbit Over 100 Million Years and Prospects for the Research in the Moon",slug:"the-evolution-of-the-moon-s-orbit-over-100-million-years-and-prospects-for-the-research-in-the-moon",totalDownloads:57,totalDimensionsCites:0,doi:"10.5772/intechopen.102392",abstract:"As a result of solving the problem of interaction of Solar-system bodies, data on the evolution of the Moon’s orbit were obtained. These data were used as the basis for the development of a mathematical model for the Moon representing its motion over an interval of 100 million years. A program of exploration of the Moon with the aim of creating a permanent base on it is outlined. Such a base is intended for exploring the Earth, the Sun, and outer space.",book:{id:"10955",title:"Lunar Science - Habitat and Humans",coverURL:"https://cdn.intechopen.com/books/images_new/10955.jpg"},signatures:"Joseph J. Smulsky"},{id:"80217",title:"Educational and Scientific Analog Space Missions",slug:"educational-and-scientific-analog-space-missions",totalDownloads:78,totalDimensionsCites:0,doi:"10.5772/intechopen.101392",abstract:"Analog space missions in Poland include international scientific, technological, and business projects designed and realized by a private research company Analog Astronaut Training Center Ltd. (AATC) devoted to the future Moon and Mars exploration. Growing experience in educational aspect of the training as well as continuous development of the habitat and its professional space science laboratory equipment correspond to increased interest of educational organizations, universities, and individual students. We serve unique practical platform for space engineering, space master, and even space doctoral theses. In addition to a wide range of training courses offered for future astronauts, for example, diving, skydiving, rocket workshops, and stratospheric missions, AATC provides a private laboratory to simulate the space environment. It carries out scientific experiments focused on biology and space medicine, as well as addressing several multidisciplinary issues related to the Moon and Mars exploration, including space mining. The main goal of each our analog simulation is to get publishable results, what means that our analog astronauts obtain not only certification of completion of the training but also ability to continue studies and to perform it individually. This chapter summarizes methodology used by us, didactic tools, and obtained results for both educational and scientific analog simulations.",book:{id:"10955",title:"Lunar Science - Habitat and Humans",coverURL:"https://cdn.intechopen.com/books/images_new/10955.jpg"},signatures:"Agata Maria Kołodziejczyk and M. Harasymczuk"},{id:"79544",title:"Regolith and Radiation: The Cosmic Battle",slug:"regolith-and-radiation-the-cosmic-battle",totalDownloads:113,totalDimensionsCites:0,doi:"10.5772/intechopen.101437",abstract:"This chapter discusses regolith utilization in habitat construction mainly from the point of view of radiation protection of humans on missions of long duration. It also considers other key properties such as structural robustness, thermal insulation, and micrometeoroid protection that all have to be considered in parallel when proposing regolith-based solutions. The biological hazards of radiation exposure on the Moon are presented and put in the context of lunar exploration-type missions and current astronaut career dose limits. These factors guide the research in radiation protection done with lunar regolith simulants, which are used in research and development activities on Earth due to the reduced accessibility of returned lunar samples. The ways in which regolith can be used in construction influence its protective properties. Areal density, which plays a key role in the radiation shielding capacity of a given material, can be optimized through different regolith processing techniques. At the same time, density will also affect other important properties of the construction, e.g. thermal insulation. A comprehensive picture of regolith utilization in habitat walls is drawn for the reader to understand the main aspects that are considered in habitat design and construction while maintaining the main focus on radiation protection.",book:{id:"10955",title:"Lunar Science - Habitat and Humans",coverURL:"https://cdn.intechopen.com/books/images_new/10955.jpg"},signatures:"Yulia Akisheva, Yves Gourinat, Nicolas Foray and Aidan Cowley"}],onlineFirstChaptersTotal:5},preDownload:{success:null,errors:{}},subscriptionForm:{success:null,errors:{}},aboutIntechopen:{},privacyPolicy:{},peerReviewing:{},howOpenAccessPublishingWithIntechopenWorks:{},sponsorshipBooks:{sponsorshipBooks:[],offset:0,limit:8,total:null},allSeries:{pteSeriesList:[{id:"14",title:"Artificial Intelligence",numberOfPublishedBooks:9,numberOfPublishedChapters:89,numberOfOpenTopics:6,numberOfUpcomingTopics:0,issn:"2633-1403",doi:"10.5772/intechopen.79920",isOpenForSubmission:!0},{id:"7",title:"Biomedical Engineering",numberOfPublishedBooks:12,numberOfPublishedChapters:104,numberOfOpenTopics:3,numberOfUpcomingTopics:0,issn:"2631-5343",doi:"10.5772/intechopen.71985",isOpenForSubmission:!0}],lsSeriesList:[{id:"11",title:"Biochemistry",numberOfPublishedBooks:32,numberOfPublishedChapters:318,numberOfOpenTopics:4,numberOfUpcomingTopics:0,issn:"2632-0983",doi:"10.5772/intechopen.72877",isOpenForSubmission:!0},{id:"25",title:"Environmental Sciences",numberOfPublishedBooks:1,numberOfPublishedChapters:12,numberOfOpenTopics:4,numberOfUpcomingTopics:0,issn:"2754-6713",doi:"10.5772/intechopen.100362",isOpenForSubmission:!0},{id:"10",title:"Physiology",numberOfPublishedBooks:11,numberOfPublishedChapters:141,numberOfOpenTopics:4,numberOfUpcomingTopics:0,issn:"2631-8261",doi:"10.5772/intechopen.72796",isOpenForSubmission:!0}],hsSeriesList:[{id:"3",title:"Dentistry",numberOfPublishedBooks:8,numberOfPublishedChapters:129,numberOfOpenTopics:2,numberOfUpcomingTopics:0,issn:"2631-6218",doi:"10.5772/intechopen.71199",isOpenForSubmission:!0},{id:"6",title:"Infectious Diseases",numberOfPublishedBooks:13,numberOfPublishedChapters:113,numberOfOpenTopics:3,numberOfUpcomingTopics:1,issn:"2631-6188",doi:"10.5772/intechopen.71852",isOpenForSubmission:!0},{id:"13",title:"Veterinary Medicine and Science",numberOfPublishedBooks:11,numberOfPublishedChapters:105,numberOfOpenTopics:3,numberOfUpcomingTopics:0,issn:"2632-0517",doi:"10.5772/intechopen.73681",isOpenForSubmission:!0}],sshSeriesList:[{id:"22",title:"Business, Management and Economics",numberOfPublishedBooks:1,numberOfPublishedChapters:19,numberOfOpenTopics:2,numberOfUpcomingTopics:1,issn:"2753-894X",doi:"10.5772/intechopen.100359",isOpenForSubmission:!0},{id:"23",title:"Education and Human Development",numberOfPublishedBooks:0,numberOfPublishedChapters:5,numberOfOpenTopics:1,numberOfUpcomingTopics:1,issn:null,doi:"10.5772/intechopen.100360",isOpenForSubmission:!0},{id:"24",title:"Sustainable Development",numberOfPublishedBooks:0,numberOfPublishedChapters:15,numberOfOpenTopics:5,numberOfUpcomingTopics:0,issn:null,doi:"10.5772/intechopen.100361",isOpenForSubmission:!0}],testimonialsList:[{id:"13",text:"The collaboration with and support of the technical staff of IntechOpen is fantastic. The whole process of submitting an article and editing of the submitted article goes extremely smooth and fast, the number of reads and downloads of chapters is high, and the contributions are also frequently cited.",author:{id:"55578",name:"Antonio",surname:"Jurado-Navas",institutionString:null,profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002bRisIQAS/Profile_Picture_1626166543950",slug:"antonio-jurado-navas",institution:{id:"720",name:"University of Malaga",country:{id:null,name:"Spain"}}}},{id:"6",text:"It is great to work with the IntechOpen to produce a worthwhile collection of research that also becomes a great educational resource and guide for future research endeavors.",author:{id:"259298",name:"Edward",surname:"Narayan",institutionString:null,profilePictureURL:"https://mts.intechopen.com/storage/users/259298/images/system/259298.jpeg",slug:"edward-narayan",institution:{id:"3",name:"University of Queensland",country:{id:null,name:"Australia"}}}}]},series:{item:{id:"24",title:"Sustainable Development",doi:"10.5772/intechopen.100361",issn:null,scope:"
\r\n\tTransforming our World: the 2030 Agenda for Sustainable Development endorsed by United Nations and 193 Member States, came into effect on Jan 1, 2016, to guide decision making and actions to the year 2030 and beyond. Central to this Agenda are 17 Goals, 169 associated targets and over 230 indicators that are reviewed annually. The vision envisaged in the implementation of the SDGs is centered on the five Ps: People, Planet, Prosperity, Peace and Partnership. This call for renewed focused efforts ensure we have a safe and healthy planet for current and future generations.
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\r\n\tThis Series focuses on covering research and applied research involving the five Ps through the following topics:
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\r\n\t1. Sustainable Economy and Fair Society that relates to SDG 1 on No Poverty, SDG 2 on Zero Hunger, SDG 8 on Decent Work and Economic Growth, SDG 10 on Reduced Inequalities, SDG 12 on Responsible Consumption and Production, and SDG 17 Partnership for the Goals
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\r\n\t2. Health and Wellbeing focusing on SDG 3 on Good Health and Wellbeing and SDG 6 on Clean Water and Sanitation
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\r\n\t3. Inclusivity and Social Equality involving SDG 4 on Quality Education, SDG 5 on Gender Equality, and SDG 16 on Peace, Justice and Strong Institutions
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\r\n\t4. Climate Change and Environmental Sustainability comprising SDG 13 on Climate Action, SDG 14 on Life Below Water, and SDG 15 on Life on Land
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\r\n\t5. Urban Planning and Environmental Management embracing SDG 7 on Affordable Clean Energy, SDG 9 on Industry, Innovation and Infrastructure, and SDG 11 on Sustainable Cities and Communities.
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\r\n\tThe series also seeks to support the use of cross cutting SDGs, as many of the goals listed above, targets and indicators are all interconnected to impact our lives and the decisions we make on a daily basis, making them impossible to tie to a single topic.
",coverUrl:"https://cdn.intechopen.com/series/covers/24.jpg",latestPublicationDate:"June 28th, 2022",hasOnlineFirst:!0,numberOfPublishedBooks:0,editor:{id:"262440",title:"Prof.",name:"Usha",middleName:null,surname:"Iyer-Raniga",slug:"usha-iyer-raniga",fullName:"Usha Iyer-Raniga",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002bRYSXQA4/Profile_Picture_2022-02-28T13:55:36.jpeg",biography:"Usha Iyer-Raniga is a professor in the School of Property and Construction Management at RMIT University. Usha co-leads the One Planet Network’s Sustainable Buildings and Construction Programme (SBC), a United Nations 10 Year Framework of Programmes on Sustainable Consumption and Production (UN 10FYP SCP) aligned with Sustainable Development Goal 12. The work also directly impacts SDG 11 on Sustainable Cities and Communities. She completed her undergraduate degree as an architect before obtaining her Masters degree from Canada and her Doctorate in Australia. Usha has been a keynote speaker as well as an invited speaker at national and international conferences, seminars and workshops. Her teaching experience includes teaching in Asian countries. She has advised Austrade, APEC, national, state and local governments. She serves as a reviewer and a member of the scientific committee for national and international refereed journals and refereed conferences. She is on the editorial board for refereed journals and has worked on Special Issues. Usha has served and continues to serve on the Boards of several not-for-profit organisations and she has also served as panel judge for a number of awards including the Premiers Sustainability Award in Victoria and the International Green Gown Awards. Usha has published over 100 publications, including research and consulting reports. Her publications cover a wide range of scientific and technical research publications that include edited books, book chapters, refereed journals, refereed conference papers and reports for local, state and federal government clients. She has also produced podcasts for various organisations and participated in media interviews. She has received state, national and international funding worth over USD $25 million. Usha has been awarded the Quarterly Franklin Membership by London Journals Press (UK). Her biography has been included in the Marquis Who's Who in the World® 2018, 2016 (33rd Edition), along with approximately 55,000 of the most accomplished men and women from around the world, including luminaries as U.N. Secretary-General Ban Ki-moon. 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\r\n\tThe demographic shifts are creating interesting challenges. People are living longer, resulting to an aging demographic. We have a large population of older workers and retirees who are living longer lives, combined with a declining birthrate in most parts of the world. Businesses of all types are looking at how technology is affecting their operations. Several questions arise, such as: How is technology changing what we do? How is it transforming us internally, how is it influencing our clients and our business strategy? It is about leveraging technology to improve efficiency, connect with customers more effectively, and drive innovation. The majority of innovative companies are technology-driven businesses. Realizing digital transformation is today’s top issue and will remain so for the next five years. Improving organizational agility, expanding portfolios of products and services, creating, and maintaining a culture of innovation, and developing next -generation leaders were also identified as top challenges in terms of both current and future issues.
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\r\n\tThe most sustained profitable growth occurs when a company expands its core business into an adjacent space. This has significant implications for management because innovation in business ecosystems differs from traditional, vertically integrated firms. Every organization in the ecosystem must be aware of the bigger picture. Innovation in ecosystems necessitates collaborative action to invent and appraise, efficient, cross-organizational knowledge flows, modular architectures, and good stewardship of legacy systems. It is built on multiple, interconnected platforms. Environmental factors have already had a significant impact in the West and will continue to have an impact globally. Businesses must take into account the environmental impact of their daily operations. The advantage of this market is that it is expected to grow more rapidly than the overall economy. Another significant challenge is preparing the next generation of leaders to elevate this to the number one priority within the next five years. There can be no culture of innovation unless there is diverse leadership or development of the next generation of leaders; and these diverse, next-generation leaders are the ones who will truly understand the digital strategies that will drive digital transformation.
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