Studies in plants that directly correlate metabolite accumulation to increased salinity stress tolerance.
\\n\\n
Released this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\\n\\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
Note: Edited in March 2021
\\n"}]',published:!0,mainMedia:{caption:"Highly Cited",originalUrl:"/media/original/117"}},components:[{type:"htmlEditorComponent",content:'IntechOpen is proud to announce that 191 of our authors have made the Clarivate™ Highly Cited Researchers List for 2020, ranking them among the top 1% most-cited.
\n\nThroughout the years, the list has named a total of 261 IntechOpen authors as Highly Cited. Of those researchers, 69 have been featured on the list multiple times.
\n\n\n\nReleased this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\n\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
Note: Edited in March 2021
\n'}],latestNews:[{slug:"webinar-introduction-to-open-science-wednesday-18-may-1-pm-cest-20220518",title:"Webinar: Introduction to Open Science | Wednesday 18 May, 1 PM CEST"},{slug:"step-in-the-right-direction-intechopen-launches-a-portfolio-of-open-science-journals-20220414",title:"Step in the Right Direction: IntechOpen Launches a Portfolio of Open Science Journals"},{slug:"let-s-meet-at-london-book-fair-5-7-april-2022-olympia-london-20220321",title:"Let’s meet at London Book Fair, 5-7 April 2022, Olympia London"},{slug:"50-books-published-as-part-of-intechopen-and-knowledge-unlatched-ku-collaboration-20220316",title:"50 Books published as part of IntechOpen and Knowledge Unlatched (KU) Collaboration"},{slug:"intechopen-joins-the-united-nations-sustainable-development-goals-publishers-compact-20221702",title:"IntechOpen joins the United Nations Sustainable Development Goals Publishers Compact"},{slug:"intechopen-signs-exclusive-representation-agreement-with-lsr-libros-servicios-y-representaciones-s-a-de-c-v-20211123",title:"IntechOpen Signs Exclusive Representation Agreement with LSR Libros Servicios y Representaciones S.A. de C.V"},{slug:"intechopen-expands-partnership-with-research4life-20211110",title:"IntechOpen Expands Partnership with Research4Life"},{slug:"introducing-intechopen-book-series-a-new-publishing-format-for-oa-books-20210915",title:"Introducing IntechOpen Book Series - A New Publishing Format for OA Books"}]},book:{item:{type:"book",id:"9356",leadTitle:null,fullTitle:"European Local Pig Breeds - Diversity and Performance. A study of project TREASURE",title:"European Local Pig Breeds - Diversity and Performance",subtitle:"A study of project TREASURE",reviewType:"peer-reviewed",abstract:'Local or autochthonous pig breeds represent a pool of genetic diversity of porcine species and a link with old-style traditional production systems and traditional pork products. These breeds were largely abandoned because they were not competitive in the concept of modern, industrial type of pig production. Despite an increased interest for local pig breeds in the past years, they remain largely untapped and the knowledge about their characteristics is limited, which was a challenge undertook in the project TREASURE in the frame of multicriteria evaluation of local pig breeds. The book represents a valuable compendium of data on census, breeding organisations, production systems, and performances with ambition to present their contemporary (preserved) phenotype.\r\nThis work has received funding from the European Union’s Horizon 2020 research and innovation programme under grant agreement No 634476 for project with acronym TREASURE. The content of this book reflects only the authors\' view and the European Union Agency is not responsible for any use that may be made of the information it contains.
\r\n\tThe adverse effects of flood disasters in urban areas have been increasing in severity and extent over the past years. The volume of losses resulting from these events is also growing exponentially, particularly in highly urbanized urban areas, where the effects of intensive land use and climate change are particularly extreme. All this despite our scientific knowledge, technical competence, and computational capacity to develop highly sophisticated and accurate forecasting and simulation models being higher than ever. In order to tackle this global issue, it is fundamental to keep on promoting and developing fundamental and applied research that allows the better targeting of interventions to improve resilience, reduce vulnerability and enhance recovery, as well as assist decision-makers in delivering more effective flood risk-reduction policies. This book will aim at contributing to this goal by gathering recent studies and state-of-the-art methodologies focused on understanding existing and emerging flood risk drivers in a climate change context.
",isbn:"978-1-80356-603-0",printIsbn:"978-1-80356-602-3",pdfIsbn:"978-1-80356-604-7",doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!1,isSalesforceBook:!1,isNomenclature:!1,hash:"c829bdd1a2a84b4b2c31ce5eaab865e2",bookSignature:"Dr. Tiago Miguel Ferreira and Associate Prof. Haiyun Shi",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/11487.jpg",keywords:"Flood Risk, Flood Hazard, Climate Change, Extreme Events, Precipitation Extremes, Extreme Events Statistics, Forecasting, Machine Learning, Statistical Hydrology, Urban Flood Risk, Urban Flood Management, Heritage Buildings",numberOfDownloads:52,numberOfWosCitations:0,numberOfCrossrefCitations:0,numberOfDimensionsCitations:0,numberOfTotalCitations:0,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"March 16th 2022",dateEndSecondStepPublish:"April 13th 2022",dateEndThirdStepPublish:"June 12th 2022",dateEndFourthStepPublish:"August 31st 2022",dateEndFifthStepPublish:"October 30th 2022",dateConfirmationOfParticipation:null,remainingDaysToSecondStep:"3 months",secondStepPassed:!0,areRegistrationsClosed:!0,currentStepOfPublishingProcess:4,editedByType:null,kuFlag:!1,biosketch:"Dr. Ferreira is ranked among the top 2% of scientists in the world in building and construction, with a long track record of publications in the field of urban risk management. Recipient of several awards and over £11 Million attracted in research grants. He has ten years of academic experience in the conservation of historical structures and vulnerability assessment under flood, earthquake, and fire.",coeditorOneBiosketch:"Dr.Shi is a pioneering researcher in the field of hydrology and water resources, mainly focusing on the topics of hydrological extremes (floods and droughts) under climate change. He has served as the main convener of two sessions in the AOGS (Asia Oceania Geosciences Society) annual meeting and the main convener of one session in the AGU (American Geophysical Union) Fall meeting.",coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"450075",title:null,name:"Tiago Miguel",middleName:null,surname:"Ferreira",slug:"tiago-miguel-ferreira",fullName:"Tiago Miguel Ferreira",profilePictureURL:"https://mts.intechopen.com/storage/users/450075/images/system/450075.jpg",biography:"Dr. Tiago Miguel Ferreira is an Assistant Professor in Civil Engineering at the University of the West of England, United Kingdom, and an invited Assistant Professor at the University of Coimbra, Portugal. Currently, he is an Editor-in-Chief of Conservar Património, a SCOPUS and Web-of-Science indexed journal dedicated to heritage studies, and a Co-Editor-in-Chief of GeoHazards, a multidisciplinary open-access journal devoted to theoretical and applied research across the whole spectrum of geomorphological hazards, namely endogenous and exogenous hazards, and those related to climate change and human activity. Ten years of academic experience in the conservation of historical structures and vulnerability assessment under flood, earthquake, and fire. 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His research interests include digital watersheds, hydroinformatics, climate change, hydrological extremes (floods and droughts), and sustainable development. He has been invited as the reviewer of over 30 international journals and the Expert Reviewer of the IPCC Sixth Assessment Report (Working Group II). He has served as the Editorial Board Member/Guest Editor of 5 international journals. 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Globally, soil salinity affects 6–10% of the total land surface (~ 800 million ha), 20–33% of which is used for agricultural purposes [1, 2, 3]. The damages caused by soil salinity are more notable in arid and semiarid regions where limited rainfall, high evapotranspiration rates, and extreme temperatures coupled with poor water and soil management practices exacerbate this problem [4, 5]. Despite the current agricultural problems due to soil salinity, world population growth has exerted substantial pressure for increased crop production to meet the global demand for food. Simultaneously, climate change has continued to markedly challenge the growth and production of agricultural crops due to variations in temperature, shifts in precipitation, reduced solar radiation, and increased evaporative demand.
Salinization can be the result of natural causes, such as flooding, wind erosion, and the redistribution of salts in shallow groundwater systems due to the weathering of minerals and salty rocks that release sodium chloride (NaCl) and other soluble salts like magnesium, calcium, sulfates (SO4), and carbonates (CO3) into the environment [6, 7]. Other sources of salinity include high evaporation rates in the tropics and the precipitation of ocean salt by wind and rain, which increase ion concentrations in soils [8]. However, the global increase in salinization is mainly due to anthropogenic activities, such as irrigation, land clearing, deforestation, agricultural intensification, the use of imbalanced fertilizers, or poor drainage systems [7]. Irrigation practices have also elevated groundwater levels, resulting in a subsequent increase in evaporation. Moreover, industrial wastewater and effluents enriched in salts can elevate the salinity levels of agricultural soils.
Salinity has drastic consequences in plants at a physiological level because it restricts the ability of plants to take up water. In plants under salt-stress conditions, the rates of transpiration and photosynthesis decrease while the opening and closing of stomata is generally quick; in addition, ion toxicity, membrane instability, and mineral limitation are observed along with the inhibition of both enzymes and metabolic pathways [9, 10, 11]. Additional problems emerge when excessive amounts of salts enter a plant and reach toxic levels, impairing normal germination, plant growth, lateral bud development, productivity, and senescence [12]. The overall results of salinity stress can be seen in impaired plant growth, physiological functions, and crop yields, although these depend on the severity of the stress, the time scale of the response, and on whether the stress was abruptly or gradually imposed [11].
Plant damage caused by salinity primarily takes place in two phases. First, increases in soil salt content cause an osmotic effect, which reduces the soil water potential and consequently plant water absorption. Second, the excessive uptake of ions, mainly Na+, Cl-, and SO2, interferes with various metabolic processes in plants, including photosynthesis, nitrogen assimilation, malate metabolism, and protein translation [13]. This nutrient imbalance reduces the amounts of calcium, magnesium, and potassium in the plant [14]. In addition, oxidative damage is generated as a result of an increase in ROS due to a reduction in the rate of photosynthesis. The combination of these factors negatively affects plant functions, including those related to metabolism, and may damage plant structures, which can ultimately lead to plant death [15, 16].
The ability of crops to grow and reproduce in saline soil is dependent on the developmental stage of the plant and greatly differs between species, which is mainly due to the variability associated with limiting salt uptake from the soil and effectively compartmentalizing it at the cellular level [17]. To deal with the adverse effects of salinity, plants trigger different physiological and biochemical mechanisms that allow them to survive and grow, including salt exclusion, the control of ion uptake and translocation, ion compartmentalization within different cells and tissues, nutrient ion transport, the synthesis of compatible solutes and osmoprotectants, morphological and anatomical modifications, membrane and hormone changes, and antioxidative metabolism responses based on the production of antioxidant enzymes and compounds [8, 13, 18]. This chapter is focused on presenting the principal determinants of salt-stress tolerance in plants, which include the antioxidant defense system, enzymatic and nonenzymatic compounds, and the most important metabolites involved in osmotic adjustments.
The antioxidant defense system is a vital mechanism by which plants deal with oxidative stress under saline conditions [19, 20, 21]. Salinization reduces the rate of photosynthesis, which alters the electron flow from central transport chains to oxygen-reduction pathways, leading to an overproduction of ROS, such as superoxide radicals (O2•−), singlet oxygen (O2), hydrogen peroxide (H2O2), hydroxyl radicals (OH•), and alkoxy radicals (RO•). ROS are generated in the cytosol, chloroplasts, mitochondria, and apoplastic spaces of cells [22, 23, 24, 25]. Although ROS are formed in biological systems as by-products of respiration and photosynthesis, in low concentrations, they act as signal transduction molecules that are involved in mediating programmed cell death, development, and responses to pathogen infections and environmental stressors [23, 26]. However, an excess of ROS produces phytotoxic reactions, biomolecule oxidation, cell membrane damage, protein degradation, enzyme inhibition, and DNA mutations [27, 28, 29]. Therefore, fluctuations in the properties and functions of ROS will eventually lead to metabolic and physiological problems in plants.
The overproduction of ROS imbalances homeostasis at cellular and subcellular levels and may ultimately lead to cellular death [30]. The intensity of the oxidative stress depends on the types of ROS produced, their concentrations, the sites where they are released, interactions with other cellular molecules, and the developmental stage and potential of the cell [31]. To avoid damage caused by the overproduction of ROS, plants employ an antioxidant defense mechanism that contains several enzymatic and nonenzymatic constituents that mitigate potential negative effects by converting ROS into less toxic molecules [32]. Given that increases in antioxidant levels usually improve plant tolerance, an adjustable control system that balances ROS production and scavenging via antioxidant enzyme and nonenzyme activities is essential for limiting toxicity levels in plants under adverse conditions [33, 34].
Plants rely on enzymes that are specifically involved in ROS detoxification, namely those that act to lower ROS levels or avoid oxidative stress, including superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX), and other enzymes involved in the AsA-GSH cycle, such as glutathione reductase (GR), monodehydroascorbate reductase (MDHAR), and dehydroascorbate reductase (DHAR). These antioxidant enzymes are located in different sites within plant cells and work together to detoxify ROS [35]. In addition, low molecular mass antioxidant compounds like ascorbate, glutathione, flavonoids, tocopherols, and carotenoids are also crucial for ROS homeostasis in plants [36, 37, 38].
Superoxide dismutase, which shows great variation across different plant species, is a metalloenzyme that plays an important role in combating oxidative stress in all subcellular organelles sensitive to ROS (i.e., chloroplasts, mitochondria, peroxisomes). This metalloenzyme catalyzes the dismutation of O2• − into O2 and H2O2. A particularly interesting property of SODs is that they can only react with O2•−) through diffusion and electrostatic guidance at limited rates, yet they are highly effective at removing O2•− [39]. SOD enzymes can be classified into three types according to their metal cofactors: copper/zinc (Cu/Zn-SOD; found in the cytosol or in plastids), manganese (Mn-SOD; found in mitochondria and peroxisomes), or iron (Fe-SOD; found in chloroplasts). Of note, it is only plants that have all three SOD types [39]. Superoxide dismutases are considered the first line of defense against ROS because an increase in SOD activity has been frequently linked to improved plant tolerance in the presence of environmental stress [40]. For example, the overproduction of SOD has been found to enhance salt-stress tolerance in sweet potatoes [40]. Moreover, SOD activity has been put forth as an indirect means to screen for plants that are salinity resistant.
Catalase was the first enzyme to be identified with antioxidant capabilities and is mainly found in peroxisomes and glyoxysomes. The function of CAT is to convert H2O2 into H20 and O2 [41]. H2O2 has been found to be produced in cell organelles during photorespiratory oxidation, the β-oxidation of fatty acids, purine catabolism, and because of the activities of enzymes, such as xanthine oxidase and SOD [42]. CAT plays a dynamic role in eliminating H2O2 by degrading it in an energetically efficient manner. For example, one CAT molecule can transform about 6 million H2O2 molecules into H2O and O2 in roughly one minute [43, 44].
Whereas animals contain a single CAT gene, plants have a multigene family that includes multiple isoenzymes (depending on the species) whose expression is regulated according to how they are distributed within tissues or organs and the environmental conditions present [44]. In plants, there are three classes of catalases that are determined based on location: class I (photosynthetic tissues), class II (vascular tissues), and class III (seeds and young seedlings) [45]. A concomitant increase in CAT activity has been observed as part of the antioxidant defense system in plants. This increase may be a manifestation of the adaptive responses of plants to abiotic stress. It may also be assumed that without any concomitant increase in CAT activity, plant growth is likely to be severely limited. Therefore, different environmental stressors constitute major drivers that either enhance or restrict CAT activity, depending on the intensity, duration, and type of stress [46].
The Asa-GSH cycle, also known as the Halliwell–Asada cycle, is a series of coupled redox reactions involving four enzymes: APX, MDHAR, DHAR, and GR [47]. This cycle plays a crucial role in the antioxidant protection system in the presence of H2O2, which is generated in different cellular compartments like chloroplasts, mitochondria, peroxisomes, and the cytosol and apoplast [48, 49]. H2O2 is scavenged by APX via the oxidation of ascorbate. This enzyme comprises a family of five APX isoforms that are located in several organelles and cellular compartments, including glyoxysomes (gmAPX), thylakoids (tAPX), the cytosol (cAPX), mitochondria (mAPX), and soluble chloroplast stroma (sAPX) [46]. APX uses two molecules of ascorbate to reduce H2O2 to H2O and produces two molecules of monodehydroascorbate (MDHA) from the oxidation of ascorbate. MHDA is converted back to ascorbate by MDHAR, which is a flavin adenine dinucleotide (FAD) enzyme with high specificity for MHDA. Subsequently, MDHA is further rapidly converted to dehydroascorbate (DHA) by the monomeric thiol enzyme DHAR. DHAR uses reduced glutathione (GSH) as the reducing substrate, which is regenerated by GR from its oxidized form glutathione disulfide (GSSG), and NADPH molecules act as donors of reducing equivalents [29, 43]. Recent studies have shown that these enzymes become active and participate in the ascorbic acid-glutathione cycle under salt-stress conditions, indicating that their increased activity may improve plant tolerance [50, 51, 52]. Figure 1 shows the functions of the antioxidant enzymes described above.
The orange squares show ROS scavenging by the antioxidant enzymes superoxide dismutase (SOD) and catalase (CAT) and the enzymes involved in the Asa-GSH cycle, namely ascorbate peroxidase (APX), monodehydroascorbate (MDHAR), dehydroascorbate reductase (DHAR), and glutathione reductase (GR). The purple circles show the nonenzymatic antioxidant compounds glutathione and ascorbate that are described in section 2.2. The diagram was modified from [
Ascorbic acid (well known as Vitamin C) is the main antioxidant in plants and acts as a major redox buffer and cofactor for enzymes. The structure of ascorbic acid is like those of hexose sugars and is comprised of a conjugated structure composed of a five-carbon lactone ring, containing enediol groups on C2 and C3. Ascorbic acid is considered to be the most powerful ROS scavenger due to its ability to donate electrons to a wide range of electron receptors in several enzymatic and nonenzymatic reactions [53]. Ascorbic acid protects cellular membranes by directly scavenging O2• − and OH while acting as a cofactor for violaxanthin deepoxidase, sustaining the dissipation of excess excitation energy in chloroplasts [54]. In addition, ascorbic acid has the redox potential to interact with hydroxyl radicals,, superoxides, oxidized glutathione, and tocopherol radicals [53, 55].
Ascorbic acid is involved in many plant pathways related to photosynthesis, hormone biosynthesis, antioxidant regeneration, defense responses, signal transduction, flowering, cell division, and growth and is abundantly found in meristems and photosynthetic cells [56, 57]. Under normal physiological conditions, ascorbic acid largely remains in a reduced form in leaves and chloroplasts [55] and accumulates in photosynthetic organs, although it can be found in high concentrations in non-photosynthetic tissues with the amounts varying depending on the plant species, variety, tissue type, and growth environment [53, 58].
Multiple experiments have been developed to evaluate the roles of antioxidant compounds, such as glutathione and ascorbic acid, when they are applied to different plant tissues. For example, Aliniaeifard et al. [59] sprayed 2 mM AsA and 3 mM GSH on olive plants treated with 100 mM NaCl and reported that plant growth parameters significantly improved with the application of AsA when compared with those of the plants sprayed with GSH. In addition, the Na + and Cl– concentrations decreased in the olive plants sprayed with AsA while the K+ concentration and K/Na ratio increased. Aliniaeifard et al. concluded that the exogenous application of AsA is recommended for improving the tolerance of olive plants under saline conditions. Similarly, Sadak [60] applied AsA to flax cultivars irrigated with salt water and found that AsA allowed for an increase in antioxidant defense via the activation of ROS scavenging enzymes.
Together with ascorbic acid, glutathione is one of the most important nonenzymatic antioxidants. Glutathione is a thiol tripeptide composed of γ-glutamyl-cysteinylglycine and plays a central role in antioxidant defense by scavenging ROS, thereby maintaining redox homeostasis in plant tissues [61]. The chemical reactivity, relative stability, and high water solubility of the thiol group of glutathione makes it ideal for protecting plants against oxidative stress or stress due to heavy metals and exogenous or endogenous organic chemicals [62]. Glutathione is localized in all cell compartments, such as the chloroplasts, cytosol, vacuoles, mitochondria, and endoplasmic reticulum. In physiological processes, glutathione plays important roles in xenobiotic detoxification, metabolite conjugation, signal transduction, and stress-responsive gene expression [61, 63]. Glutathione also participates in tissue growth and development, cell death and senescence, and the enzymatic regulation of pathogen resistance [64].
Glutathione also appears to be important in controlling O2•−, •OH, and H2O2 levels in cells under stressful conditions given the observed changes in the ratio of its reduced (GSH) and oxidized (GSSG) forms, which provides information on the cellular redox balance [65, 66]. GSH is required for GRX-mediated dithiol and monothiol reduction mechanisms and is important for the activation of GPX, GST, and glyoxalase functions, particularly recycling GSH from GSSG in the presence of NADPH. Therefore, GSH plays a key role in ROS tolerance under oxidative stress in association with other ROS-metabolizing enzymes [57]. In addition, the primary reaction in plants when they are exposed to progressive salinity is the oxidation of GSH, indicating that this reaction is an early response to oxidative stress [33, 67].
Studies have demonstrated the benefits associated with the application of GSH in modulating salt-stress tolerance and positively influencing yield-contributing traits in plants like rice (
Flavonoids are largely responsible for the pigmentation of seeds, flowers, stems, roots, and fruits. Flavonoids are aromatic compounds that are derived from chalcone, which is obtained from phenylalanine and malonyl-coenzyme A. According to their chemical structures, flavonoids can be classified into flavones, flavanones, flavonols, flavans, flavandiols, isoflavones, anthocyanins, and condensed tannins [70, 71].
Flavonoids act to prevent ROS increase through the inhibition of ROS-generating enzymes, including cyclooxygenase, lipoxygenase, monooxygenase, and xanthine oxidase. The ability to chelate metals is essential for impeding the regeneration of radical ROS molecules, such as superoxide, hydroxyl radicals, and hydrogen peroxide [72, 73]. Given that they contain functional hydroxyl groups, flavonoids constitute a secondary antioxidant system, as they support the function of other ROS scavenging systems when the activities of antioxidant enzymes are reduced. In addition, flavonoids serve as antioxidant barriers that protect cellular components against oxidizing pollutants like ozone (O3) and sulfur dioxide (SO2). Due to their elevated abilities to donate electrons or hydrogen atoms, the antioxidant capacities of flavonoids have been found to be many times higher than those of either ascorbic acid or a-tocopherol [74, 75].
Interestingly, in
Tocopherols constitute a group of lipophilic compounds that are important antioxidant and nonenzymatic components capable of inhibiting lipid peroxidation. Tocopherols are also essential for maintaining membrane integrity. These lipophilic compounds are synthesized from homogentisic acid and isopentenyl diphosphate in the plastid envelope. Tocopherol exists in four isomeric forms (α-, β-, γ-, and δ-), with the predominant forms being α- and γ-tocopherol. The composition of tocopherol depends on the genotypic features of the plant, growth conditions, tissue type, and stress intensity. Tocopherols are present in seeds, fruits, roots, tubers, cotyledons, hypocotyls, stems, leaves, and flowers. In addition, α-tocopherol may regulate the concentrations of plant hormones, such as jasmonic acid, which control both growth and development [38, 77].
As a component of thylakoid membranes, tocopherol acts as an important scavenger of 1O2 and OH while maintaining a stable redox status and the structure and function of PSII. Tocopherols have also been found to reduce lipid peroxyl radicals (obtained from lipid peroxidation) to their corresponding hydroperoxides [78]. In addition, tocopherols participate in cell signaling and they may protect embryos during germination from ROS, senescence, and stress [79, 80]. Recent studies have also shown that tocopherol plays a positive role under salt-stress conditions by controlling Na+/K+ homeostasis and the hormonal balance while minimizing oxidative stress [81]. Additionally, it has been demonstrated that the foliar application of a-tocopherol in onion plants and
Carotenoids are hydrophobic pigments derived from isoprene that play essential roles in photosynthesis and nutrition and protect against photo-oxidative damage in higher plants [37]. Carotenoids are mainly present in the form of 40-carbon tetraterpene, which consists of eight isoprene units [84]. In plants, carotenoids are synthesized in plastids from isoprenoid precursors. Lycopene is a principal carotenoid and is a product of the sequential desaturations of phytoene and may be converted into b-carotene by lycopene b-cyclase (Lcy-b) [85, 86]. Carotenoids act as light harvesters by dissipating excess energy as heat, and they also protect the photosynthetic apparatus from the free radicals produced during photosynthesis by stabilizing photosystem I and II and the thylakoid membrane. Furthermore, carotenoids are precursors of hormones like abscisic acid and strigolactones [87].
Carotenoids quench the O2 and H2O2 formed under salt stress due to their low triplet state energy and in this way, protect lipids from peroxidation while suppressing radical chain reactions [88, 89, 90]. Li et al. [91] recently showed that the overexpression of three genes associated with carotenoid biosynthesis from
Salinity decreases the osmotic potential of soil, which in turn limits water absorption by plants. In addition, the toxicity of Na + and Cl- impedes the uptake of essential nutrients. In particular, Na + toxicity can disrupt the absorption of K+ and Ca2+, favoring the production of ROS and inducing oxidative damage [92].
Plants have developed responses to deal with water loss and the transport and accumulation of toxic ions. Given that water loss in plants originates due to the low osmotic potential of the soil, plants must employ compensatory mechanisms [93]. Plant cells permit higher influxes of Na+, which decrease the water potential and cause osmotic stress in cells. Salts are translocated from the roots to shoots to maintain ion homeostasis, although this reduces photosynthetic efficiency due to the inhibition of photosystem II (PSII) [94]. To cope with osmotic stress and maintain ion homeostasis, plants biosynthesize osmolytes.
Osmolytes or compatible solutes are small molecules with low molecular weights that are electrically neutral, highly soluble, and do not affect normal biochemical processes [95]. The biosynthesis and accumulation of osmolytes constitutes an essential protection strategy for plants under abiotic stress. Multiple studies have elucidated the roles that osmolytes play in abiotic stress tolerance due to natural accumulation [96, 97], exogenous application [98, 99], or the transgenic expression of osmolyte pathway genes [100, 101], which are summarized in Table 1.
Proline | Exogenous application | [102, 103, 104] | |
GABA | Natural accumulation | [105, 106] | |
Putrescine | Natural accumulation | [107] | |
Spermidine | Exogenous application | [108] | |
Spermine | Natural accumulation | [109] | |
Glycine betaine | Exogenous application | [110, 111, 112] | |
Glucose and fructose | Natural accumulation | [113, 114] | |
Trehalose | Transgenic expression | [115] | |
Raffinose | Transgenic expression | [116] | |
Mannitol | Transgenic expression | [117] | |
Myo-inositol and pinitol | Transgenic expression | [118, 119] | |
GABA = Gamma amino butyric acid |
Studies in plants that directly correlate metabolite accumulation to increased salinity stress tolerance.
The primary function of osmolytes is to accumulate inside the cell to maintain the osmotic balance between the plant cell and its surroundings [120]. Osmolytes can also act as chemical chaperones by stabilizing and protecting proteins and membranes [121] or by acting as ROS scavengers [122]. Some of these compatible solutes are highly effective in reducing the extent of K+ loss in response to salt stress [123]. Overall, osmolytes are chemically diverse, although they can be broadly grouped into amino acids, polyamines, betaines, sugars, and polyols, which are shown in Figure 2.
Chemical structures of the most common osmolytes observed in salt-stress responses in plants. These structures can be grouped into amino acids, polyamines, betaines, sugars, and polyols.
Amino acids are the building blocks of proteins, which carry out many biological processes and provide structure and support for cells [124]. In addition, many studies have reported the accumulation of amino acids in plants exposed to drought and salinity conditions [97, 98, 125, 126]. This accumulation may be due to elevated amino acid production and/or the stress-induced breakdown of proteins [127]. Some of the amino acids that have been found to accumulate are alanine, arginine, glycine, serine, leucine, valine, and proline, in addition to non-protein amino acids like citrulline, ornithine, and gamma amino butyric acid (GABA) [128, 129].
Proline is the most important osmolyte and signaling molecule and generally accumulates in the cytosol. Proline also acts as an ROS scavenger and molecular chaperone, stabilizing the structure of proteins and protecting cells from potential damage induced by osmotic and oxidative stress [120, 130, 131]. Proline is mainly synthesized from glutamate by Δ1-pyrroline-5-carboxylate (P5C) synthetase and P5C reductase enzymes, although under nitrogen-limited conditions, proline is synthesized in the ornithine pathway and transformed into P5C/GSA (glutamate-1-semialdehyde) via ornithine-δ-aminotransferase [132, 133].
Proline has been associated with increased salt tolerance in tomato (
Gamma amino butyric acid is a four-carbon non-protein amino acid with an amino group attached to γ-carbon instead of α- carbon. GABA is mainly synthesized from glutamate in the cytosol by glutamate decarboxylase and then transported to the mitochondria [136]. GABA metabolism has been associated with the carbon/nitrogen balance and ROS scavenging [137, 138]. In addition, GABA accumulation and metabolism have been found to be activated by salt exposure in tobacco plants [105]. Moreover, GABA-T
Polyamines (PAs) are low molecular weight aliphatic nitrogenous bases containing two or more amino groups [140]. The most common PAs in higher plants are putrescine, spermidine, and spermine. Putrescine can be produced from ornithine by ornithine decarboxylase or from arginine by arginine decarboxylase. Putrescine is then converted to spermidine (spermidine synthase) and spermidine into spermine (spermine synthase). Both spermidine and spermine can be converted back to putrescine by polyamine-oxidases [36].
PAs regulate diverse cellular functions that are essential for cell growth, including senescence, development, cell proliferation, and signal transduction while also regulating the expression of genes in response to various stressors. However, PA accumulation has been detected in plants under abiotic stress [141]. Polyamines act as osmolytes due to their ability to block ion channels (cationic structures) and scavenge ROS [36]. Exogenous spermidine treatment in both salt-sensitive and salt-tolerant rice cultivars has been found to result in plasma membrane recovery after injury induced by salinity [108]. In addition, the exogenous application of spermidine and putrescine was found to increase the postharvest shelf-life of
Glycine betaine (GB) is a quaternary ammonium compound whose distribution among plants is restricted to certain species like
The exogenous application of GB mitigates the adverse effects of salinity stress in some plant species. For example, the foliar application of GB was found to result in significantly improved salt tolerance in rice plants [110], and the exogenous application of GB on tomato plants subjected to salt stress resulted in an increase in fruit yield of ~40% compared with that of untreated plants [149]. Salinity tolerance in response to GB has also been observed in mung bean (
Sugars provide carbon and energy for cellular metabolic processes while regulating plant growth and development. However, under stressful conditions, carbohydrate metabolism results in an increase in sugar levels. The production and collection of soluble sugars directly contributes to ROS scavenging, osmotic adjustments, carbon storage, and the stabilization of protein structures like Ribulose-1,5-bisphosphate carboxylase–oxygenase [151]. Sugars are also the main osmolytes that participate in osmotic adjustments and can contribute up to 50% of the total osmotic potential in some plant species [152]. Structurally, all kinds of sugars have been reported in response to salinity, including monosaccharides (glucose and fructose), disaccharides (trehalose and sucrose), and oligosaccharides (raffinose family) (Table 1).
Studies have shown that plants mobilize starch and fructans from storage organs (roots, stems, and amyloplasts in leaves) to increase the accumulation of sugars, such as glucose, fructose, and sucrose. Starch hydrolysis by the b-amolytic pathway represents the primary means of starch degradation in leaves under normal growth conditions and may also be involved in stress-induced starch hydrolysis. Downton [113] shows that
Similarly, trehalose accumulates in many organisms due to various abiotic stressors and has been reported to act as an osmolyte [154]. Trehalose is a disaccharide that is synthesized by a two-step process in which trehalose-6-phosphate is first synthesized from glucose-6-phosphate and then dephosphorylated to trehalose by trehalose-6-phosphate phosphatase [155]. Several studies have evaluated the transgenic expression of trehalose biosynthesis genes, which have been shown to enhance trehalose metabolism and tolerance to abiotic stress. For instance, the overexpression of otsA and otsB in transgenic rice plants was found to result in the increased accumulation of trehalose content and an overall improved photosynthetic capacity, reducing oxidative damage and improving ion uptake and partitioning under conditions of salt stress [115]. Also, the induction of OsTPP1 and OsTPP2 by the exogenous application of ABA was found to enhance cold, salinity, and drought tolerance in rice plants [156].
The raffinose family oligosaccharides (RFO), such as raffinose, stachyose, and verbascose, are macromolecules that act to protect plants from drought, salt, cold, freezing, and oxidative stress [157]. In addition, RFOs have been implicated in membrane protection and radical scavenging. Furthermore, Arabidopsis plants overexpressing Arabidopsis GolS1 or GolS2 were found to accumulate high levels of galactinol and raffinose and were more tolerant to salinity stress [116].
Polyols or sugar alcohols are polyhydric alcohols that are widely distributed in the plant kingdom, and they can be grouped into linear and cyclic structures. The most common polyols are mannitol, glycerol, sorbitol (linear), and myo-inositol, along with the methylated derivatives ononitol and pinitol (cyclic) [158, 159]. Polyols accumulate in the cytoplasm and act as osmoprotectants and ROS scavengers, preventing water loss and oxidative damage to membranes and enzymes. Polyols originate via the reduction of aldoses or their phosphate esters and are generally water soluble in nature [160].
Williamson [161] reported that mannitol, sorbitol, and inositol increase drought and salinity tolerance in some plants. Transgenic tobacco plants overexpressing bacterial mannitol-1-phosphate dehydrogenase were found to accumulate high concentrations of mannitol in their leaves and roots and exhibited a high degree of salt tolerance [101]. Moreover, Adams et al. [162] reported that myo-inositol, ononitol, and pinitol accumulated under salt-stress conditions in several halotolerant plant species. Furthermore, the over-expression of MIPS and IMT from halotolerant plants was found to increase cyclic polyol levels and salt-stress tolerance in tobacco plants [118, 119].
Salt stress is one of the most important factors that limits the productivity of agricultural crops and threatens global food security. Salinity produces osmotic stress and ionic toxicity in plants, which alters cell homeostasis and reduces plant water absorption, adversely affecting growth and plant productivity. Osmotic imbalances have been found to cause an overproduction of ROS that leads to the oxidation of biomolecules, cell membrane damage, protein degradation, enzyme inhibition, DNA mutations, and cellular death. To limit the damage, cells turn on enzymatic and nonenzymatic antioxidant machinery. The ROS detoxification system is very complex and controlled at multiple levels because ROS participate in biological processes like photosynthesis, signal transduction, development, and programmed cell death. Therefore, to evaluate the negative and positive roles of ROS, it is important to understand tolerance mechanisms in plants and the ability to control or moderate ROS levels. Recently, it has been proposed that ROS homeostasis may be modulated through the use of nanoparticles that activate the expression of antioxidant enzyme genes or that have the ability to scavenge ROS [22], thus improving salt tolerance.
Another response to salinity in plants is the induction of osmolyte biosynthesis to stave off water loss and the accumulation of toxic ions. The production and accumulation of osmolytes inside the cell is essential for the protection and survival of plants under salt stress and other forms of environmental stress, although their production is taxonomically restricted. It has been shown that the exogenous application of osmolytes, such as aminoacids, betaines, or sugars, and the overexpression of osmolyte genes constitute viable options to improve plant resistance to salinity. Thus, the use and application of osmolytes in agricultural settings should continue to expand. Current studies have shown that when taken together, metabolic and physiological plant responses can provide valuable information of the possible mechanisms required for plants to adapt to stress. Thus, modulating one component of the antioxidative defense system or a particular metabolite might be insufficient to confer resistance to the entire plant. To generate salt-tolerant species, it is essential to use modified plants that incorporate more than one component, such as the use of multiple antioxidants or metabolites or a combination of both. In this way, further progress in genomics, proteomics, and metabolomics, together with improvements in biotechnological tools, will allow for the underlying mechanisms of salt-tolerant species to be fully elucidated and understood.
We would like to thank Programa para el Desarrollo Profesional Docente (PRODEP), University of Guadalajara, Consejo Nacional de Ciencia y Tecnología (CONACYT), and Laboratorio Nacional PlanTECC for the use of facilities and the support provided via the project 315918.
Lifestyle has changed over the world in the past decades. The industrialization process and technological advances have simplified the physical work of human beings and changed the lifestyle of the last generations. Not that long ago, most of the jobs required physical activity and some energy expenditure. Nowadays the percentage of work sectors demanding high levels of physical activity has reduced drastically. This new reality derives in many people forced to spend at more than 8 h/day sitting and having difficulties to reach the physical activity recommendations [1]. Sedentary lifestyles have become a significant public health issue spreading worldwide, although there is evidence of being linked to a range of chronic health conditions [2]. Extended periods of inactivity can produce metabolic dysfunction and impair blood sugar regulation [3], elevate blood pressure [4], and make it difficult to use fat as a metabolic substrate, as well as increase the risk of early death regardless of physical activity levels [5]. Therefore, it seems crucial to find strategies that can be applied in all life domains to be able to reduce sedentary behavior, as well as to increase physical activity. Including regular and well-structured sedentary breaks during long sitting periods could help reduce the negative effects of a sedentary lifestyle.
This chapter aimed, firstly, to provide scientific evidence of the need to reduce sedentary behaviors as well as to include regularly sedentary breaks. Secondly, to show some possibilities and examples of how to break sedentarism in daily life. We believe that introducing these practices in workspaces, schools, leisure time, and in the daily activities of older adults might help control the negative effects derived from sedentary lifestyles.
Historically, exercise physiologists have studied sedentary lifestyle as the opposite of physical activity. The terms that have been used for research in this area have been confusing, which makes it difficult to compare clinical trials. Already in the 1950s, Morris et al. [6] concluded that sedentary work increased cardiovascular risk compared to those who worked more physically active. That study, among others, resulted in a strong area of research focused, for over 60 years, on quantifying the level of physical activity necessary to reduce morbidity and mortality [6]. These investigations provided recommendations on physical activity and the implementation of public policies to promote physical activity practice.
Despite the efforts, a high percentage of the population (mostly from countries that suffered rapid urbanization and industrialization) do no reach the physical activity recommendations and the tendency is that this number increases [1] Office works, school, screen games, technology, passive transportation and sedentary leisure time have had a strong impact on reducing the opportunities to perform physical activity at the same time that promote opportunities for sedentary behavior in all the life domains.
For the past two decades, the number of studies focused on sedentary behavior has grown exponentially, and physical activity and sedentary behavior can be considered as an independent research field. The term sedentary behavior comes from Latin “sedere” which means “to sit”. But not only the “position” determines what is currently conceived as sedentary behavior. Sedentary behavior is defined as any waking behavior characterized by the expenditure of 1.5 metabolic equivalents of task (MET)s or less of energy while in a sitting, reclining, or lying posture [7]. Sedentary behavior, like physical activity, can be found in all life domains (work, study, transport, and free time). Although research in this field has increased notably in the last decade, there is still confusion in the terminology and the scientific community has not reached a consensus in some terms and concepts yet. Many definitions of sedentary behavior can be found in the literature but some common concepts are repeated, such as low energy expenditure, mostly under 1.5 METs, activities performed in sitting, lying or reclining position and while the person is awake [7]. Besides, some other concepts associated with sedentary behavior have aroused the interest of the scientific community. Sedentary bouts, breaks of sitting, sedentarism, sedentary lifestyle vs. inactivity, among others, are related terms that could help deeply understand this problem.
Sedentary Behavior Concept has suffer an evolution over the years. Although the distinction between sedentarism and physical inactivity (not meeting worldwide recommendations for physical activity) has already been settled [7], it is still common to find some confusion in terms such as sedentary time, sitting time, screen time and stationary time; which, although in some cases are overlapping concepts, refer to different behaviors [7]. Because they refer to different aspects of behavior (position, movement, effort and the use of digital implements), these traits can be blended in different ways, so that some criteria are met but others are not. For example, one can be seated but doing physical activity (cycloergometer), so it’s not a sedentary behavior; one could be stationary, but not sitting (e.g., waiting in line); one may be in sedentary behavior, but not sitting (instead lying down watching TV), which in turn is independent of screen usage (reading a book); among other examples.
Studies on bed rest [8, 9, 10, 11, 12, 13] provided useful information on the consequences of inactivity and low energy expenditure for long periods. Thanks to these studies, a lot is known about the effects of prolonged inactivity in metabolism and organ systems. Different studies focused on the effects of bed rest on metabolic function, found peripheral insulin resistance in skeletal muscle and adipose tissue, hepatic insulin resistance and a dyslipidemia [10], as well as a decline in function, muscle mass, and muscle strength [12] and a reduction in cardiorespiratory capacity after one-week bed rest [13]. In regards to the musculoskeletal structure, inactivity produces loss of strength and endurance, contractures, changes in soft tissues, disuse osteoporosis, sarcopenia, and degenerative joint disease [8]. At the cardiovascular level, the consequences can be postural hypotension, cardiac dysfunction, and thrombotic events [13]. Additionally, bed rest can lead to impaired respiratory, renal, gastrointestinal, and nervous system levels [9]. Outside hospitalization or illness, free-living healthy adults rarely spend these amounts of bed rest. Nevertheless, technological and social factors have made prolonged sitting time a common practice in all life domains (work, domestic life, and leisure time).
Measuring physical activity and sitting time is complex. Research has been aimed at improving the quality of the data through the objective measurement of sedentary behavior using accelerometry, observing that the self-report measurement underestimates the daily time of sedentary behavior concerning the objective measurement.
Researchers have focused on developing devices to be able to objectively quantify physical activity. In the past decades, many studies using accelerometers have been carried out. A multi-country study (USA, Brazil, UK, Denmark, the Czech Republic, and Hong Kong) using accelerometry found that the average sedentary time per day was 513 min/day, or 8.55 h/day [14]. Sedentary time was estimated to be responsible for 3.8% of all-cause mortality in adults according to a meta-analysis pooling data across 54 countries [15]. The United States Physical Activity Guidelines Advisory Committee (PAGAC) [16] recently comprehensively reviewed the scientific evidence, linking sedentary behavior with specific physical health indicators in adults and older adults, including mortality, cardiovascular disease, type 2 diabetes, cancer, and obesity. Moreover, high levels of sedentary behavior are also negative associated with cognitive function, depression, function and disability, physical activity levels, and health-related quality of life [17]. In contrast, little evidence has demonstrated the relationship between sedentary behavior and musculoskeletal pain, accidents or injuries, fatigue, sleep, or work productivity [18]. Ku et al. [19] published in 2018 a meta-regression analysis involving more than 1 million participants in which the cut-off points of daily sedentary time that were related to all-cause mortality in adults were established for data measured objectively and self-reported [19]. According to the results of the study, the method of measuring sitting time significantly moderated the association between daily sitting time and mortality risk. The cut-off of daily sitting time in studies with self-report data was 7 h/day in comparison with 9 h/day for those with data measured by devices.
It is accepted that exercise is an effective strategy for reducing key cardiovascular risks [20]. Nevertheless, it is unclear if the benefits can be modified by a sedentary lifestyle. Therefore, it is important to clearly define different concepts such as physical activity/inactivity or sedentary behavior, as their physiological consequences on health are different. While physical activity/inactivity is referred to whether or not a person reaches the physical activity recommendations, a person is considered as sedentary if he/she spends long periods of the day in sedentary behavior. While for the first one (cut-off points for being physically active) there is enough evidence to determine the recommendations (150 minutes of moderate physical activity or 75 minutes of vigorous physical activity or an equivalent metabolic combination between both, plus 2–3 days/week of resistance training) [21], for the second one (cut-off points for being sedentary) there are still no recommendations, since studies have found inconclusive results.
That means that a person can meet the physical activity guidelines and still be considered sedentary. Sedentary behavior might produce harmful effects on health independently of physical activity level, but when both are combined, the results seem to change (combined joint association). In other words, high levels of sedentary behavior combined with low levels of physical activity increase the risk of death by 46% [18]. On the contrary, some studies have shown that high levels of physical activity can counteract or reduce the risk of death caused by prolonged sedentary behavior [22]. Similar results were obtained in cancer patients, where in the most active patients no relationship was observed between sedentary behavior and cancer mortality, while for those less active the risk of death increased [23].
Using the concepts of sedentary and/or physically active person, we can describe four possible combinations:
Figure 1 represents graphically these possibilities.
Person’s classification according to sedentary behavior and physical activity practice. (A) Sedentary inactive, (B) non-sedentay inactive, (C) sedentary active, (D) non-sedentary active.
The health implication for possibilities A and D are clear. Classification A has a negative influence on health and is negatively associated with all-cause mortality and D is positively associated with better health markers. What is not fully clear yet, are the implications of classifications B and C. Can one the variables counteract the negative effect of too much of the other one? Or, are the positive effects of one variable suppressed by the other one?
As mentioned before, some studies found that high levels of physical activity might attenuate the increased risk of some illness or death associated with high sitting times [22]. Notwithstanding, there is still some uncertainty in the characteristics of the specific dose–response curves, which makes it difficult to determine specific quantitative public health recommendations [24]. As sedentary lifestyle in western societies does not tend to reduce, new strategies might be the solution. Some degree of sedentary lifestyle might be beneficial for health so that it helps to rest and recover. On the contrary, excessive sitting time may become a risk factor. Scientific evidence has not found an increase in the risk of death from any cause in people with a total sitting time between 4–8 hours/day when compared to those who remain seated for less than 4 hours. Nevertheless, the risk increases by 15% when sitting time rises to 8–11 hours/day, and by 40% with sitting times higher than 11 hours/day [25]. Contrary, some other studies found a dose–response relationship for every 1-hour increase in sitting time in intervals between 0–3, >3–7, and > 7 h/day total sitting and all-cause mortality. This model estimated a 34% higher mortality risk for adults sitting 10 h/day, after taking physical activity into account, although the risk increased staggered [26], similar to other studies that observed statistically significantly higher risk of death with sedentary times of 9.5 h/day or more [23].
This situation has put the focus on the double challenge of increasing levels of physical activity and reducing sedentary behavior. Many countries have developed strategies to promote changes in the population. As an example, the Canadian government created the Canadian 24-Hour Movement Guidelines for Adults (https://csepguidelines.ca/). It recommends that adults between 18–64 years must limit sedentary time to 8 hours/day or less, including no more than 3 hours/day of recreation screen time and breaking long periods of sitting as often as possible.
As it has been mentioned before, modern lifestyles predispose a high percentage of the population to spend long periods in sedentary behaviors. As too much sitting time is related to different chronic diseases such as type 2 diabetes, obesity, hypertension, and cardiovascular diseases, or some types of cancer, it seems crucial to clearly understand the mechanism and strategies to reduce the negative effects of a sedentary lifestyle. Generalizing, we get up, use the elevator to go to the car, drive to work, take the escalator to go work, spend 8 hours at least working with minimum movement, drive back home, eat, have some hours of recreational time, watch TV and go to bed. Fortunately, different lifestyles and personal situations (occupational situation and leisure-time preferences) as well as inherent individual differences, result in different accumulations of sedentary time. Due to the strong available evidence on the deleterious effects of a sedentary lifestyle on health, it is necessary to better understand the metabolic mechanisms and how it is accumulated. Researchers have observed that reducing or breaking up sedentary time may result in beneficial changes in body composition and acute improvements in markers of cardiometabolic risk.
Sedentary behavior might be considered as a multifactorial concept, where four different aspects influencing it should be taken into account:
It has been proposed that breaks in sedentary time could help counteract the negative effect of prolonged periods of whole-body inactivity. A break in sedentary time can be defined as a period of non-sedentary activity, such as standing or walking in between two sedentary bouts [7]. Experimental studies have demonstrated that interrupting sedentary time with short frequent breaks reduces daily glucose, postprandial glucose, and insulin resistance [3, 26, 27]. In a study carried out by Healy et al. [28] in 2008, the authors found, that interruptions of sedentary behavior were negatively associated with obesity and cardiometabolic health. These results highlighted, already at that time, the fact that not only total sitting matters but also how it is distributed in a period of time. The characteristic of the sedentary breaks in the study from Healy et al. showed that the breaks reported by the participants were shorter than 5 min on average, and they were performed at a light intensity. Results from this study also found lower waist circumference, BMI, triglycerides, and 2-h plasma glucose in the participants with higher sedentary break bouts, independent of total sedentary time or moderate-to-vigorous intensity activity time. Since this pioneering study was published, the scientific community have had an increased interest in analyzing the effects of sedentary breaks, to be able to deeply understand the effects of prolonged sitting on metabolism, as well as to establish clear and specific guidelines of intervention. Different types of sedentary breaks have been studied trying to analyze if shorter bouts of sitting time, are less metabolic disrupting even when the total amount of daily or weekly sitting times are similar.
Brief bouts of light-intensity-activity sedentary breaks could reduce the negative effects of long periods sitting on lower limb vascular function in healthy and overweight/obese adults [29]. Experimental studies [30, 31, 32] have seen that combining exercise with breaks in sitting resulted in additional reductions in postprandial insulin-glucose dynamics and triglycerides when comparing exercise and uninterrupted sitting. This effect, although useful in any case, seems to be more effective in those with high basal insulin resistance.
As many studies focused on analyzing the effects of sedentary breaks to counteract the metabolic problems associated with prolonged sitting time have found positive interactions, the question that remains unanswered is not if we should break sitting regularly, what already has a positive answer. The unanswered question is, which is the best structure for a sedentary break?
As it has been mentioned before, the lack of enough specific interventional studies complicates for experts to concrete the most optimal structure for sedentary breaks. A recent study by Wheeler et al. [30] investigated the effects of 3 different sitting strategies in overweight and obese: i) uninterrupted sitting for 8 h, ii) sitting for 1 h, moderate-intensity walking for 30-min and uninterrupted sitting for 6.5 h and iii) sitting for 1 h, moderate-intensity walking for 30 min and sitting for 6.5 h interrupting sitting every 30 min with 3 min of light-intensity walking. They found reductions in postprandial insulin-glucose dynamics and triglycerides by combining exercise with breaks in sitting. This study not only proposes a way to help reach the physical activity recommendation by breaking sedentary time for 30 min/day but also demonstrates that regular sedentary breaks help control the metabolic deleterious effect of prolonged sitting.
A well-controlled meta-analysis conducted by Loh et al. in 2020 [33] found that the use of sitting breaks moderately attenuated post-prandial glucose, insulin, and triacylglycerol. The authors also found that the glycemic attenuation was greater in people with a higher body mass index. An interesting result was that for attenuating glucose levels, a statistically significant small advantage for sitting breaks was found over continuous exercise when exercise matched energy. That could mean that for glucose regulation, it might be more interesting short regular breaks along the day, than one continuous bout of exercise.
The skeletal muscle might also play a key role in glycaemia control, which is even more important in overweight. Bergouignan et al. [34] performed an analysis from randomized clinical trials comparing one or three days uninterrupted sitting with sitting interrupted with light-intensity or moderate-intensity walking every 20-min in the modulation of contraction- and insulin-stimulated glucose uptake pathways in muscle. They found that both sitting break interventions reduced postprandial glucose concentration as well as a transition to modulation of the insulin-signaling pathway and increased capacity for glucose transport. The moderate-intensity intervention resulted in a greater capacity for glycogen synthesis and ATP production. These results might through some light in preventive strategy for metabolic diseases.
Published literature [35] might tend to propose that the best option to reduce the negative effects of sedentary behavior on metabolic functions could be to combine regular activity breaks of several minutes every 30 min of sitting with 30 min of continuous walking whether at the beginning or the end of the long sitting period.
Therefore, breaking sedentary time should be a good way to reduce the negative effects of long periods of sitting, for both metabolic and muscle function. These breaks are even more interesting for patients with initial high blood sugar, insulin resistance, or overweight-obesity. The general recommendation would be to make an active 2–3 min-break every 30 min of sitting time. If the activity made during these breaks is of moderate-high intensity, such as climbing stairs, the metabolic benefits might be greater.
The workplace is considered an important environment for the promotion and protection of health [36]. According to a report from the World Health Organization (WHO) together with the World Economic Forum, 65% of the world’s adult population is part of the workforce [37]. In 2007, about 3.1 billion people were part of the economically active population and it was estimated that by 2021 this number would exceed 3.6 billion [38]. Taking into account that this working adult population spends around a third of the day at work, workers´ health must be seen as a priority action.
Encouraging the reduction of sedentary behavior and promoting the practice of physical activity in the workplace is a strategy that helps maintain the health of the working population and affects their close environment. In 2018, the WHO presented the Global Action Plan for Physical Activity [39], with two mean challenges: reducing sedentary behavior by 2030 as well as the percentage of inactive population by 15% to the reported values of 2016. This plan encourages the population to take advantage of the many opportunities that arise in daily living to integrate physical activity, including the workplace (as a fundamental environment to practice physical activity programs as well as its promotion).
The activities where sedentary behavior predominates have increased lately and the workplace is a clear example. The machines have replaced human physical work at the same time that there has been a notable increase in office jobs, where the employee spends most of the working day in front of a computer. Although the negative consequences for cardiometabolic and musculoskeletal health of sedentary behavior have been widely demonstrated and office work represents for many workers a third of their day sitting, few have been made to improve this situation and reduce sitting time at workplaces, with the associated health risk.
The Healthy Work Environment model, proposed by the WHO [38] proposes intervention programs to reduce and break sedentary behavior in the workplace as a health promotion model and protection strategy. This model proposes four scenarios of action or “avenues of influence”, which are not isolated, but rather overlap each other:
The physical environment of the work, which refers to the structure, air, machinery, furniture.
The psychosocial work environment, which includes the organization of work and institutional culture, attitudes, values, beliefs that can affect the mental and physical well-being of workers.
Personal health resources in the workplace, that consist in an environment that promotes health, health services, information, resources, opportunities, and the flexibility that the companies offer to workers to support the efforts to improve or maintain healthy lifestyles, as well as to monitor and support your physical and mental health.
The physical participation of the institution in the community, which includes the activities that the company carries out to improve the safety, well-being, and quality of life of workers and their families.
To successfully establish health promotion programs in the workplace, certain conditions must be considered:
Raising awareness among managers and chiefs of the importance of these interventions, facilitating employees to carry them out. Companies’ leaders must understand that these strategies are not only not time wasted but will also result in increased productivity.
A previous evaluation of the workplace and the type of tasks that are developed, that help design an optimal plan.
Execution of the plan with the support of all interested parts (managers, middle managers, bosses, CEO, etc.) and commitment by workers.
Re-evaluation and adaptation of the proposal.
With different adaptations, similar models can be recommended with more or less the same stages.
Experts have suggested different strategies to reduce or interrupt sedentary behavior in the workplace, which could be grouped into the following categories [40].
Physical/environmental changes in workplace design
Desks with adjustable height that allow lifting them to work standing up.
Raised desks with a treadmill.
Rooms with high tables for standing meetings.
Modify the layout of the workplace, for example, by placing printers, trashcans, or water dispensers away from desks, which will force employees to stand up and walk a few steps when they need to use these items.
Provide bicycle racks, lockers, and services to wash up to encourage active transportation to work.
Eliminate architectural barriers to allow employees to move around the workplace, creating unobstructed corridors and spaces that invite walking.
Changes in workplace policy to incentivize and encourage reduction and disruption of sitting time
Promote the holding of standing or walking meetings.
Propose active breaks during working hours (short breaks in which you can do joint mobility exercises, put on a musical theme and dance, or any activity that allows interrupting the sedentary behavior through light physical activity)
Offer group physical activity practice.
Encourage the use of breaks for short walks.
Encourage employees to communicate with their colleagues by approaching their desks rather than by phone or messages.
Propose to take advantage of telephone communications to do them standing or walking (obviously, spaces that do not interfere with the work of others should be considered).
Encourage the use of the stairs instead of the elevator or escalator.
Information and advice to raise employee awareness and commitment by offering
Workshops, training courses and outreach programs on the importance of reducing sedentary behavior. Reporting on the health risks of sedentary behavior and the benefits of practicing physical activity could allow people to evaluate their behavioral choices.
Campaigns through various means, such as posters, signage, emails, WhatsApp messages, telephone calls or internal messages to motivate a change in behavior or.
Install reminder software every 30 minutes on employees’ mobile phones or personal computers, for example, to interrupt the sedentary behavior by standing up, dancing or doing some movements.
As mentioned so far, human bodies are adapted to maintain a physically active lifestyle. Proof of this is the health consequences of an insufficient level of physical activity. However, it is also true that neurobiologically we are adapted to “optimize” our energy expenditure, avoiding additional efforts when possible; In other words, sedentary behaviors are attractive for human beings, and willing power is required to counteract this attraction and opt for a behavior with higher associated energy expenditure [41]. It has been studied how the energy cost associated with a task affects, not only our decision to choose another more “economic” one, but directly to our perception of the initial task [42] and, therefore, to our future intention to undertake it.
A process as complex as human behavior cannot be reduced to just one component. Emotional/affective factors, as well as built habits, are also related to sedentary behavior and physical activity [43]. However, it is an interesting starting point if we seek an alternative approach to the one traditionally used. The assumption that human behaviors are decided by rational evaluations of the available information are underlying concepts in many current intervention strategies and, therefore, knowing the benefits of regular physical activity and the damages of prolonged sedentary behavior should be enough to solve the problem [44]. Nevertheless, in light of the sustained global pandemic of physical inactivity, it may be necessary to complement and enrich this approach with other perspectives.
Sedentary behaviors in free time are usually classified as screen-time (watching television, videos via streaming platform or physical medium, browsing the internet and social networks by both on a computer, tablet or cell phone and the use of video games) or not screen-time (sitting down to eat, participating in social gatherings, playing board games, recreational, attending cultural events such as cinema, theater, show music, sports competition, religious ceremony, doing artistic activities like writing or drawing or hobbies. The extensive list is testimony to the enormous offer of sedentary activities in free time. Recommendations on physical activity and sedentary behavior limit the amount of time in sedentary behaviors, but particularly those carried out in front of the screen [21].
Sedentary activities in front of the screen in free time, in addition to adverse effects on physical health, are related to adverse effects on mental health, mainly in minors [45]. Screen time during childhood is negatively correlated with brain connectivity, compared to time spent reading books, as well as being related to loss of imagery ability [46] or social–emotional functionality [47]. Interestingly, and in contrast to these studies, in the specific case of video games, there is evidence that indicates various cognitive benefits according to the type of game (action, strategy), and even positive socio-emotional impacts [48].
Of the large number of sedentary activities carried out in free time, although the impact on physical health is equivalent, it would be differential over other dimensions of the subject’s health. This leads to one of the perspectives mentioned in the literature as a strategy to address sedentary behavior: “harm reduction”. Assuming that certain socio-cultural (technological) changes are already part of daily life, priority is given to modifying those behaviors that present a greater health risk: replacing sedentary behaviors in front of the screen with sedentary behaviors without a screen, or by non-sedentary screen activities (for example, walking while using portable devices or replacing sedentary video games with active ones) [49].
Different classifications have been proposed for reducing sedentary behavior in the free time [50]: 1) environmental interventions such as devices that limit the time of television use), and 2) behavioral interventions like education campaigns about the harms of prolonged sedentary behavior; 3) multi-component interventions which include both types mentioned above.
Although studies on this fact do not have homogeneous methodologies, some findings can be pointed out. Studies that focus on the sedentary behavior of children in the home context have found a relationship between the existence of screen devices in the bedroom and greater sedentary behavior (with less reading time). Likewise, both the interventions that use devices that limit the use of television and those on family rules for screen use have been successful in reducing sedentary behavior. Furthermore, it was observed that in those cases in which the parents had more television time, or participated with their children in sedentary activities, the children presented higher levels of sedentary behavior. In some studies, the existence of adequate space or equipment for practicing physical activity at home is related to less sedentary behavior (although it does not present higher levels of physical activity at moderate or vigorous intensities) [51].
In the case of adults and the elderly, studies on free time are scarce and methodological imprecise. The absence of control in the domains makes it difficult to control the changes since the decrease of sedentary time in a domain does not imply its replacement by physical activity since it could simply shift to sedentary behavior in another domain. Those interventions aimed exclusively at reducing sedentary behavior have better results than those that also focused on increasing physical activity [50].
For children and adolescents, as well as for adults and the elderly, there is another alternative intervention strategy, which constitutes itself in an emerging field of research: exergaming, also known as active gaming or effort video game. These video games, unlike the traditional ones, are controlled with body movements (either full body or only certain segments); Thus, instead of being a sedentary activity, at least light-intensity physical activity is achieved (with the potential to become moderate intensity and even vigorous). In the US, it is estimated that 90% of children and adolescents play video games recreationally. In an increasing technophile society, and in which electronic entertainment is already part of our lives, exergaming stands as a strategy to address those to whom other physical activity proposals are not convincing. In addition, the commitment, immersion, and experience of “flow” that they can generate, make them a great resource for health-related purposes. Sustainability over the years of this type of activity has been investigated, finding greater adherence in women, and similar to that of team sports [52].
Results for studies analyzing experiences in exergaming as part of both school physical education and at-home context show a decrease in sedentary behavior with potential, according to the intensity at which the game is played, increase of moderate-to-vigorous physical activity and good adherence to intervention programs. One of the challenges of exergaming is the “replay value” (once the game becomes monotonous and therefore the motivation to continue playing decreases), which maintain adherence. Multiplayer games (both face-to-face and remotely) show greater adherence. The eventual increase in the number of published games would compensate for this situation, allowing simply to change to a new one [53].
Particularly interesting is the research with older people, which improvements for both institutionalized and community-dwelling subjects, and not only in the physical dimensions but also in the cognitive one [54].
In all these cases, we refer mainly to consoles-home exergaming, but everyday mobile devices with augmented reality technology (Pokémon Go with geo-location system integrated into cell phones) are great opportunities to promote exergaming. Pokémon Go requires active movement of the player around their surroundings to play. This game mechanic has achieved a statistically significant change in the number of steps per day (thus decreasing sedentary behavior), although there is still not enough evidence on long-term adherence.
In the latter case, as in some home exergaming video games, there is no explicit intention in its design to promote health effects or to prescribe a systematic physical activity program. However, they have the potential to have a positive impact on the health and well-being of those who opt for this type of digital entertainment. Sedentary behavior in free time poses a great global challenge that requires, particularly for new generations, imagination and innovative approaches, in tune with contemporary technologies and paradigms.
It is well accepted that physical activity is beneficial to maintain and improve health and well-being across life [55]. In infants, toddlers, and preschoolers, high levels of physical activity have been seen to be related to better social and motor development improved metabolic health, and decreased adiposity, while a sedentary lifestyle is related to higher adiposity and poorer psychosocial health and cognitive development [54].
Children (preschoolers and scholars) spend more than 2 h/day of screen time, which is the maximal time recommended for this age group [56], plus eating time, school, passive transportation, homework, etc., which results in more than 8 h/day of sitting at this age. Moreover, studies found that screen time was associated with an increased risk of overweight/obese independent of physical activity [54]. Sex differences were also found. Boys are generally more involved in physical activity than girls, which normally spent more time on domestic tasks and homework. Children living in rural areas tend to use more active transportation than those who live in urban areas. Older children also tend to use more active transportation than the younger ones [57]. Taking into account that sedentary behavior in children is directly associated with classical cardiovascular risk factors like elevated blood glucose levels, insulin resistance, high blood pressure, obesity, and elevated blood lipids [58], strategies that help reduce total daily sitting time in children are crucial.
Nevertheless, although childhood should be a life stage where children should freely play, run and jump as part of their natural development, social rules, obligations, parent’s overprotection, new technologies, and urban environments, hinder the practice of physical activity for children with dramatic consequences. A qualitative study performed by Hidding et al. [59] aimed in determining the reasons for children to be sitting from the children or parents perspective, found that children most repeated reason was that they sit because is the norm and they have to and because they can play better that way. Other common answers were: I sit because seated activities are fun, I sit because I’m tired, I want to relax, I want to rest, I sit because of my health, I sit because there is nobody to play with, I sit because there is nothing to do, I sit because I’m not in the mood to do anything, I sit because of the weather. In regards to the answer “I sit because there is nobody to play with”, in families with more than one child, seems to be easy for children to perform physical activity [59].
All this information brings the experts ‘awareness of the necessity of reconsider children’s environments. The CSEP Canadian 24-Hour Movement Guidelines [56] propose an Integration of physical activity (both light and moderate-to-vigorous), sedentary behavior, and sleep as the three principal parts of the day. All three must be right balanced to promote overall health, well-being, and quality of life. These guidelines use “the four S rule”:
SWEAT: Moderate to Vigorous Physical Activity: An accumulation of at least 60 min/day.
STEP: Light physical activity: Several hours of a variety of structured and unstructured light physical activities (playing, walking)
SLEEP: Uninterrupted 9 to 11 h/night for those aged 5–13 years and 8 to 10 h/night for those aged 14–17 years, with consistent bed and wake-up times.
SIT: Sedentary behavior: No more than 2 h/day of recreational screen time and limited sitting for extended periods.
Figure 2 ilustrates de cited guidelines.
Canadian 24-hour movement guidelines for children and youth (5-17 years): an integration of physical activity, sedentary behavior, and sleep. Taken from CSEP website (
In addition, parents might consider changing indoor activities for outdoor ones, when possible, and including moderate to vigorous physical activity in exchange for light physical activity at some point of the day.
Findings from a recent meta-analysis [60] on the physical activity a sedentary behavior suggest that physical activity interventions can improve adolescents’ mental health.
Worldwide, the population is aging, which results in higher economic and social costs, as well as increased numbers of people living with more health problems, as aging increases the risk of suffering from chronic diseases. Therefore, the concept of successful aging has become a priority to guarantee, not only that life expectancy is high, but also that the years lived are of the best quality possible, free or with minimum chronic diseases. Physical activity has been proven to help increase or maintain health throughout life. Due to physical activity tends to reduce with age, older adults must become a risk population. Disability, frailty, dysfunction, or sarcopenia are some of the problems that can affect older adults, which can compromise the independence level [61]. As physical activity decreases in this group, sedentary behavior increases, with fatal consequences. Maintaining physical activity levels and reducing sedentary time, should be a priority for the administrations. In this regard, there is evidence about the negative associations of sedentary behavior with frailty and how this relationship can differ by sitting bout length. Some studies [62] have found that prolonged sedentary bouts and total sedentary time were associated with higher mortality risk in frail individuals but not in robust. These results, including moderate-to-vigorous physical activity, reducing sedentary time in those frail older adults, as well as including sedentary breaks seem like a suitable strategy to prevent dependency and maintain health. As the total hours of a day are always 24, that means that when a person increases the time spent in moderate-to-vigorous physical activity, this person is reducing the time spent in another activity, that could be sitting or light physical activity. If sleeping time remains stable and a person substitutes 30 min/day of light physical activity for moderate-to-vigorous physical activity that includes resistance training, and at the same time changes 1 hour of sitting for light physical activity such as walking, the frailty status could be significantly reduced. Moreover, if this person would include a short sedentary break every 30 min - 1 hour of the total time this person is seated, the benefits would be even higher with only small changes.
Due to older adults are mostly retired, which releases them of office sitting time and have a lot of leisure time, political strategies must center on providing older adults with a safe environment where they can perform light physical activity, such as walking [63]. Pavements and sidewalks in good conditions, green areas, and safe cross-roads might help improve that older adults go more often outside to take a walk. At the same time, organized affordable exercise activities, specifically designed by experts for this population, could make that older adult reach the recommendation for moderate-to-vigorous physical activity and resistance training. Moreover, these activities also promote social interaction, which improve wellbeing and might help reduce depression and anxiety, improving health-related quality of life, as well. These two actions would help to achieve physical activity recommendations at the same time that sedentary time would be reduced. To completely promote health in this group, clinicians, governments, and media should establish campaigns to make older adults understand the importance of breaking sedentary time. Things such as get up in the commercials when they watch TV, walking or standing while they are phoning, or get up to drink some water once each hour might be enough to break sitting time.
Lifestyle has dramatically changed in the last century. Industrialization and technology have reduced the physical requirements of many jobs, urbanization has changed population habits, force them to use passive transport instead of active ones, children play with digital devices since they are very young and older adult do not have to go outside because cities, family and environment easily provide all their needs. However, this sedentary lifestyle has disastrous consequences for health. Physical activity is necessary to maintain an optimal physiological function and prolonged sitting time interferes with the proper metabolic regulation. The combination of both, low physical activity levels and prolonged sitting time, maybe even more deleterious. That suggests a double challenge for developed countries; reducing and stopping prolonged sedentary behavior as well as increasing levels of physical activity. Although each of them separately has concrete effects on health, their interaction must be also taken into account. Sedentary behavior appears to be negative for health “per se”, as well as low physical activity levels, but how both are combined is what can make the difference. Scientific evidence says that high physical activity levels might help counteract the negative effects of sitting time and that this effect is progressive. That means, that the higher the physical activity intensity, the less negative effects of sitting time. At the same time, it seems that long continuous sitting bouts are more harmful than the same total sitting time but with breaks in between. Eight hours seated without any break might be a lot worse for metabolic regulation than the same 8 h of sitting but with breaks of 2–3 min every 30 min-1 h. With all these ideas in mind, the strategy to reduce sedentary behavior seems clear: practice enough physical activity, reduce free-time sitting and screen time, promote active transportation, and include sedentary breaks at sedentary jobs. The reason why these strategies are not working is complex and implies a compromise at different levels. First, governments must provide opportunities for affordable exercise practice and physical activity-friendly environments. Secondly, at workplaces, managers, CEOs, and bosses must be aware of the importance of promoting working places where employees have the opportunity of breaking sedentary time, and that it is seen as normal. Third, citizens should make efforts to include active activities in the free time as well as substitute classic videogames for exergaming, where at least, sitting time is exchanged for light physical activity. Last but not least, special populations (children and older adults) should not be forgotten. Parents and schools should reconsider the rules and norms and adapt them, when possible, to others more active versions, not forcing children to spend long periods seated promoting at the same time at least one hour of physical activity per day. Controlling screen time and giving good examples must be another priority for parents. In regards to older adults, societies should allow them to perform easy tasks that increase physical activity, encouraging them to used active transportations to carry them out, at the same time that exercise programs, specifical design for this population, are easily available in every neighborhood.
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\n\nThe first step in obtaining funds for your Open Access publication begins with your institution or library. IntechOpen’s publishing standards align with most institutional funding programs. Our advice is to petition your institution for help in financing your Open Access publication.
\n\nHowever, as Open Access becomes a more commonly used publishing option for the dissemination of scientific and scholarly content, in addition to institutions, there are a growing number of funders who allow the use of grants for covering OA publication costs, or have established separate funds for the same purpose.
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