Dimensions of the internet parenting style (adapted from [8], p. 89).
\r\n\tAnimal food additives are products used in animal nutrition for purposes of improving the quality of feed or to improve the animal’s performance and health. Other additives can be used to enhance digestibility or even flavour of feed materials. In addition, feed additives are known which improve the quality of compound feed production; consequently e.g. they improve the quality of the granulated mixed diet.
\r\n\r\n\tGenerally feed additives could be divided into five groups:
\r\n\t1.Technological additives which influence the technological aspects of the diet to improve its handling or hygiene characteristics.
\r\n\t2. Sensory additives which improve the palatability of a diet by stimulating appetite, usually through the effect these products have on the flavour or colour.
\r\n\t3. Nutritional additives, such additives are specific nutrient(s) required by the animal for optimal production.
\r\n\t4.Zootechnical additives which improve the nutrient status of the animal, not by providing specific nutrients, but by enabling more efficient use of the nutrients present in the diet, in other words, it increases the efficiency of production.
\r\n\t5. In poultry nutrition: Coccidiostats and Histomonostats which widely used to control intestinal health of poultry through direct effects on the parasitic organism concerned.
\r\n\tThe aim of the book is to present the impact of the most important feed additives on the animal production, to demonstrate their mode of action, to show their effect on intermediate metabolism and heath status of livestock and to suggest how to use the different feed additives in animal nutrition to produce high quality and safety animal origin foodstuffs for human consumer.
",isbn:"978-1-83969-404-2",printIsbn:"978-1-83969-403-5",pdfIsbn:"978-1-83969-405-9",doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!1,hash:"8ffe43a82ac48b309abc3632bbf3efd0",bookSignature:"Prof. László Babinszky",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/10496.jpg",keywords:"Technological Feed Additives, Feed Industry, Quality of Compound Feed, Non-Antibiotic Growth Promoter, Product Quality, Additive Enzymes, Digestibility of Nutrients, NSP Enzymes, Farm Animals, Livestock, Immunity, Microbiome",numberOfDownloads:null,numberOfWosCitations:0,numberOfCrossrefCitations:null,numberOfDimensionsCitations:null,numberOfTotalCitations:null,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"November 24th 2020",dateEndSecondStepPublish:"December 22nd 2020",dateEndThirdStepPublish:"February 20th 2021",dateEndFourthStepPublish:"May 11th 2021",dateEndFifthStepPublish:"July 10th 2021",remainingDaysToSecondStep:"2 months",secondStepPassed:!0,currentStepOfPublishingProcess:4,editedByType:null,kuFlag:!1,biosketch:"Professor Emeritus from the University of Debrecen, Hungary who authored 297 publications (papers, book chapters) and edited 3 books. Member of various committees and chairman of the World Conference of Innovative Animal Nutrition and Feeding (WIANF).",coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"53998",title:"Prof.",name:"László",middleName:null,surname:"Babinszky",slug:"laszlo-babinszky",fullName:"László Babinszky",profilePictureURL:"https://mts.intechopen.com/storage/users/53998/images/system/53998.jpg",biography:"László Babinszky is Professor Emeritus of animal nutrition at the University of Debrecen, Hungary. From 1984 to 1985 he worked at the Agricultural University in Wageningen and in the Institute for Livestock Feeding and Nutrition in Lelystad (the Netherlands). He also worked at the Agricultural University of Vienna in the Institute for Animal Breeding and Nutrition (Austria) and in the Oscar Kellner Research Institute in Rostock (Germany). From 1988 to 1992, he worked in the Department of Animal Nutrition (Agricultural University in Wageningen). In 1992 he obtained a PhD degree in animal nutrition from the University of Wageningen.He has authored 297 publications (papers, book chapters). He edited 3 books and 14 international conference proceedings. His total number of citation is 407. \r\nHe is member of various committees e.g.: American Society of Animal Science (ASAS, USA); the editorial board of the Acta Agriculturae Scandinavica, Section A- Animal Science (Norway); KRMIVA, Journal of Animal Nutrition (Croatia), Austin Food Sciences (NJ, USA), E-Cronicon Nutrition (UK), SciTz Nutrition and Food Science (DE, USA), Journal of Medical Chemistry and Toxicology (NJ, USA), Current Research in Food Technology and Nutritional Sciences (USA). 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Venkateswarlu",coverURL:"https://cdn.intechopen.com/books/images_new/371.jpg",editedByType:"Edited by",editors:[{id:"58592",title:"Dr.",name:"Arun",surname:"Shanker",slug:"arun-shanker",fullName:"Arun Shanker"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"878",title:"Phytochemicals",subtitle:"A Global Perspective of Their Role in Nutrition and Health",isOpenForSubmission:!1,hash:"ec77671f63975ef2d16192897deb6835",slug:"phytochemicals-a-global-perspective-of-their-role-in-nutrition-and-health",bookSignature:"Venketeshwer Rao",coverURL:"https://cdn.intechopen.com/books/images_new/878.jpg",editedByType:"Edited by",editors:[{id:"82663",title:"Dr.",name:"Venketeshwer",surname:"Rao",slug:"venketeshwer-rao",fullName:"Venketeshwer Rao"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"4816",title:"Face Recognition",subtitle:null,isOpenForSubmission:!1,hash:"146063b5359146b7718ea86bad47c8eb",slug:"face_recognition",bookSignature:"Kresimir Delac and Mislav Grgic",coverURL:"https://cdn.intechopen.com/books/images_new/4816.jpg",editedByType:"Edited by",editors:[{id:"528",title:"Dr.",name:"Kresimir",surname:"Delac",slug:"kresimir-delac",fullName:"Kresimir Delac"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"69756",title:"Molecular Tools for Gene Analysis in Fission Yeast",doi:"10.5772/intechopen.84896",slug:"molecular-tools-for-gene-analysis-in-fission-yeast",body:'Schizosaccharomyces pombe (S. pombe) is a single-cell, nonpathogenic yeast, described in Germany in 1893 by P. Linder, named “pombe,” and was originally isolated from East African millet beer [1]. S. pombe is an ascomycete fungus, whose lineage is evolutionarily remote from the yeast Saccharomyces cerevisiae [2]. Actually, S. pombe is phylogenetically as distant from budding yeasts as it is from humans. In 1950, two homothallic strains, h90 (968) and h40, and two heterothallic strains with opposite mating types, h− and h+, were isolated [3, 4]. There are several heterothallic strains with different genomic configurations at the mating type locus, but the heterothallic strains commonly used in the laboratory are h+N (975) and h−S (972) [5].
S. pombe is also called fission yeast because it is divided by binary fission. However, it has two forms of reproduction: one by binary fission and another by sporulation. Therefore, it is possible to find it in both haploid and diploid states. S. pombe cells are cylindrical, 3–4 μm in diameter and 7–15 μm long in haploid state, while in diploid state, they measure from 4 to 5 μm in diameter and 20–25 μm long [6]. S. pombe was the sixth eukaryote to have its entire genome sequenced [7]. The genome of S. pombe has a size of 13.8 Mb and is organized in chromosome I of 5.7 Mb, chromosome II of 4.6 Mb, and chromosome III of 3.5 Mb, along with a mitochondrial genome of 20 Kb [8]. It contains the ribosomal RNA genes 5.8S, 18S, and 25S with a length of approximately 1.1 Mb [9]. Approximately, 4940 genes encoding proteins (including 11 mitochondrial genes) and 33 pseudogenes have been predicted. Almost 50% of fission yeast genes have at least one intron, and in total, there are 5300 introns in 2510 protein-coding genes. The process of splicing also appears to be more similar to splicing in human cells (
In its haploid state and in favorable conditions, S. pombe grows through a mitotic cycle. The optimal growth temperature for S. pombe cells is 30°C (25–36°C) with a doubling time of 2–4 hours [11]. In both haploid and diploid cells, S. pombe mitotic cell cycle is organized into the G1, S, G2, and M phases. There are two major controls regulating progress through the cell cycle: the G1–S transition and the G2–M transition. Both points are regulated by cyclin-dependent serine/threonine protein kinase Cdc2 [12, 13]. The meiotic cell cycle is a modified mitotic (M) cell cycle [14, 15]. Like other eukaryotes, in the meiotic cell cycle of fission yeast, MI is a reductional division, without intervening S phase before the second meiotic division (MII). The meiosis process concludes with the formation of an ascus including four haploid spores [15].
Under conditions of nutrient restriction especially nitrogen, cells become arrested in G1, and if the two sexual types (h+ and h−) are present, they will conjugate to form a diploid zygote, known as zygotic ascus [16, 17]. In a similar way as in mammals, two cells of opposite sex are recognized by the system of communication of pheromones. A cell experiment a polarized morphogenesis in the direction in the direction of the pheromones source cell by a process called shmooing. Next, two cells will merge by conjugation or “mating” producing a zygote. The zygote is diploid and could be kept in a diploid mitotic cycle if the conditions of the medium improve at this point in the cycle. If the growing conditions are unfavorable, the diploid yeast will enter into meiosis to culminate with the formation of an ascus with four haploid spores. The spores will germinate and enter again into the mitotic cycle when the environmental conditions allow it, thus closing the cycle.
S. pombe has become one of the best-studied eukaryotes today. Dr. Forsburg gave it the name of micromammal [18]. In fission yeast, genes and proteins homologous to higher eukaryotes have been described related to recombination, chromosomal organization, chromatin modification, stress response mechanisms, DNA damage response, mitosis, meiosis, cell cycle control, mRNA splicing, cell morphogenesis and polarity, and post-translational modifications of proteins such as glycosylation [19, 20, 21, 22, 23, 24, 25, 26, 27].
In fission yeast, gene deletion or one-step gene deletion by gene replacement via homologous recombination is probably the most used molecular tool in the functional characterization of the function of the gene and the protein. Gene disruption is a genetic analysis strategy to achieve gene modifications, generation of tagged protein fusions, genetic expression placed under the control of a regulated promoter, specific mutations, insertions, and deletion [28, 29]. Gene replacement by homologous recombination in S. pombe has allowed the construction of chromosomal interruptions of genes such as sts1, gcs1, gsh2, hmt1 [30], and git2/cyr1 [31].
Gene replacement requires a switch construction that contains 5′ and 3′ homologous regions of the target locus that flank a selection marker gene, and its efficiency in homologous recombination depends largely on the size of these regions [28]. The genetic construct is incorporated into the cells by transformation, then the reporter gene that will be used for gene replacement is inserted into the target gene due to the presence of terminal homologous regions in the construct, thus eliminating a large fragment of the target gene and incorporating instead, the reporter gene. At the beginning of the use of this technique, the protocol was based on obtaining the homologous regions to the target gene by digestion to flank a selection marker gene that was obtained from a plasmid containing the desired selection gene as well as the restriction sites resulting from the digestion of the homologous regions.
For this method, it was essential that DNA fragments share the restriction site for subsequent linkage. With the advance in molecular biology, methods based on PCR were developed [32]. The PCR strategy was improved by Wang et al. [32], and the protocol described the generation of construction switch gene called two-step PCR. Four oligonucleotides are required for the amplification of the homologous regions of the target gene to eliminate or modify. These PCR fragments can be called AB located in the 5′ region and CD located in the 3′ region of the target gene. A novel strategy was a little modification in the 3′ antisense oligonucleotide from AB region and 5′ sense oligonucleotide from CD region, which contains a short complementary sequence and a single restriction site to facilitate the link of the two products generated in the first PCR and then forming a product that serves as template for the second PCR. The final PCR product can be called ABCD and is cloned into a plasmid. At the same time, the gene marker selection used in the gene replacement like leu1 or ura4+ is amplified by PCR with oligonucleotides including the same restriction site in both ends, used in the 3′ antisense oligonucleotide from AB region and 5′ sense oligonucleotide from CD region of target gene to replacement. The PCR product of marker gene is incorporated into a cloning vector, and then it’s digested with the unique restriction enzyme selected. Finally, the plasmid containing ABCD fragment is digested with the same restriction enzyme used to prepare the marker gene and linked to produce the AB-selection marker gene-CD gene deletion cassette.
In order to achieve the gene modification, a one-step gene deletion technique by pop-in homologous must be performed [29]. The gene deletion cassette is transformed into a yeast strain with a deletion in the endogenous gene selected like ura+ (ura4-D18) to the gene replacement [33] by the lithium acetate protocol [34]. Then, it is efficiently targeted to its homologous location in the chromosome DNA. Moreover, it is widely known that the efficiency of homologous recombination is greatly stimulated if the incoming DNA sequence has free ends. The DNA flanking to the marker gene, on each side, recombines with the genome, inserting the marker gene into the target gene, therefore disrupting or completely replacing it.
It has been reported that the optimal length of homologous sequences to achieve an efficient elimination of the gene is 80–100 pb. Nevertheless, high efficiency in mutagenesis directed for S. pombe has been brought, using long segments of homologia of the gene target (≥250 pb), with efficiencies in the homologous integration of up to 100% [35]. The selection marker genes used in S. pombe are based on gene markers of Saccharomyces cerevisiae. The most used genes are ura3, leu2, ade6, trp1, and his3 that synthesize enzymes used for the biosynthesis of uracil, leucine, adenine, tryptophan, and histidine, respectively [36]. A high and efficient integration in the strains that have mutations in the locus leu1-32 and Ura4-294 of S. pombe with own genes leu1+ and ura4+ has been showed.
In addition, to make the functional analyses of various genes as well as minimize incidental recombination events between DNA sequences within the marker gene and a chromosomal sequence, gene deletion cassettes consisting entirely of heterologous DNA sequences have been designated. Those gene deletion cassettes even allow multiple gene deletions to be performed. Because the incorporation of loxP sites flanking the marker gene allows Cre recombinase-mediated rescue, the marker can be reused for the next gene deletion. Genes can be deleted in sequential order using different gene deletion cassettes carrying different selectable markers. Then, a gene deletion cassette would be removed from the chromosome DNA by mitotic or recombinase-mediated recombination. The strategy allows the use of the recyclable deletion cassettes, useful to disrupt the next gene of interest with the same marker gene [37, 38].
Fission yeast is a very popular system for protein expression with potential biotechnological applications. The choice of yeasts for the purification of proteins, their structural analysis, and the generation of mutants aimed at knowing the function of proteins is based on the shared conserved biological processes as cell cycle progression, protein turnover, vesicular trafficking, and signal transduction with cells of higher eukaryotes [39, 40]. In yeasts, the appropriate expression of proteins with the posttranslational modifications required allows to obtain the correct protein structure and function. So, the use of yeast in the industrial production of enzymes employed in food, medicine and health, environment, and other applications has been proposed [41, 42]. To fulfill this purpose, “humanized yeast model systems” have been created as tools to study the molecular mechanisms involved in chronic degenerative diseases such as neurological disorders [43, 44]. Due to the accessibility of the yeast to simple genetic and environmental manipulations, it reduced complexity compared to the mammalian models.
Fission yeast is an excellent system to study the complex intracellular mechanisms underlying neurodegenerative diseases such as Alzheimer’s disease (AD). Heterologous expression of Tau and Aβ can provide new insights into the pathobiology of these proteins in vivo as well as the screening of compounds that may be useful in treatment and/or prevention of AD [45]. Recently, it was reported that ginger (dietary condiment) fermented with S. pombe had neuroprotective effects on in vivo models of AD. FG improved recognition memory, ameliorated memory impairment in amyloid beta1–42 (Aβ1–42) plaque-injected mice, reinstated the pre- and postsynaptic protein levels decreased by amyloid plaque toxicity, as well as attenuated memory impairment in Aβ1–42 plaque-induced AD mice [46].
Numerous expression vectors have been used in molecular studies on S. pombe [47, 48, 49]. A typical plasmid of S. pombe contains an origin of bacterial replication, an antibiotic resistance gene to select recombinant cells in bacteria, an autonomous replication sequence (ARS1), and a marker of selection of yeast. More complex plasmids can include a regulated or constitutive promoter, a transcription terminator, or epitope tags [47, 50].
The use of antibiotics to induce genes to antibiotic resistance genes as selection markers into the yeast plasmid is very frequent. The kanamycin/G418, hygromycin B, phleomycin/bleomycin, and nourseothricin/clonNat are excellent markers in fission yeast [51]. Relative to auxotrophy, new markers such as ade7, his1, his2, his3, his5, arg3, arg12, lys1, lys2, and tyr1 are being developed [51, 52, 53, 54, 55].
However, ade6, his3+, LEU2, and ura4+ remain the most widely used markers for the selection of multi-copy vectors in common use. The pDUAL series and pJK148 vectors have been used to achieve the conversion of the leucine auxotrophy of leu1.32 to leucine prototrophy to select integration at the leu1 locus by recombination as well as pJK210 has been used to rescue ura4.294 to target integration at the ura4 locus [36].
In regard to the promoters used in the cloning vectors to protein expression, there are many promoters between the most used such as adh1+, which is a constitutive promoter. The fbp1+ is repressed by exogenous cAMP. The SV40 promoter is of constitutive expression. The CaMV promoter is tetracycline inducible. The inv1+ is glucose repressible. The ctr4+ is copper repressible and nmt1 (strong, intermediate, and weak promoters) is thiamine repressible [47]. The latter is the most used promoter and was the first characterized in the expression of protein heterologous. The nmt1 (no message in thiamine) promoter (Pnmt1) is considered as an inducible/repressible strong promoter that directs the transcription. It can be repressed by the addition of thiamine to a medium or induced in the absence of thiamine [56]. Pnmt1 has excellent dynamic range and a low off-state transcription but takes 14–16 h to induce upon thiamine withdrawal. Pnmt1 responds to the lack of exogenous thiamine and is induced approximately 75-fold when thiamine is removed from the growth media. However, the activity of Pnmt1 is repressed by the yeast extract present in a medium rich in YE and YES. So, some modifications into the TATA box of Pnmt1 have been made. Variants of this promoter were developed to reduce both off-state and on-state transcription [57, 58]. Pnmt4 and Pnmt8 are excellent options to choose the desired level of expression. However, an induction of transcription of 14–16 his also required.
To solve the problem, other promoters were generated to avoid the inactivation of the promoter nmt1 in the YES culture medium. The promoters of the 276-bp eno and 273-bp gpd were modified from eno101 and gpd3 genes in S. pombe. Both are stronger and constitutive promoters, which increase 1.5-fold higher expression of lacZ gene than nmt1 promoter. In addition, the 276-bp eno and 273-bp gpd promoters were not affected by the components of YES medium like Pnmt1.
As it was mentioned, there are other constitutive promoters widely used in S. pombe. The CaMV 35S promoter is a moderate constitutive promoter in S. pombe derived from the native 35S promoter of the plant viral cauliflower mosaic virus through deletion of the Tet repressor [59]. The adh1+ promoter of alcohol dehydrogenase is constitutively transcribed at high levels in cells grown in glucose and glycerol. However, the adh1+ promoter is weaker than the nmt1 promoter and may only be useful if a low level of gene expression is desired [58, 60]. Padh1 has two mutant variants, namely Padh41 (a mildly weak version of the adh1+ promoter) and Padh81 (a weak version of the adh1+ promoter, where the TATA box sequence TATAAATA is changed into TA), and both of these promoters express the downstream gene constitutively. Padh81 has been used in the study of the dynamic of the kinetochores [61, 62].
Therefore, it is necessary to find more efficient promoters for high-expression proteins in S. pombe. Other induction systems have rapid response times, but have a short dynamic range or relatively high levels of off-state transcription. The lsd90 promoter that is strongly induced by heat stress was cloned into the pJH5 vector, which contains an ARS element and a truncated URA3m as selectable marker. Following the expression of the luciferase reporter into the vector and making the comparison with other promoters such as Pnmt1, Padh1, and AOX1, it was found that lsd90 promoter promotes a constitutive expression of luciferase, at a level of 19-, 39-, and 10-fold higher than the promoters above mentioned, respectively [63]. The urg1 gene was identified as a rapidly induced transcript, responding to uracil addition in ~30 min and exhibiting low off-state transcription and high dynamic range [64] Other useful constitutive promoters in the protein expression are tif471 (with moderate force) and lys7 (weak promoter) [27, 65].
The pREP series vectors are general-purpose episomal vectors widely used in fission yeast research that contains a replication origin ARS1, ura4+, or LEU2 as the selective marker and kan, nat, hph, and bsd genes as a second type of marker of resistance to the specific antibiotics G418, clonNAT, hygromycin B, and blasticidin S, respectively. The latter are used routinely during chromosomal integration. The pREP vectors have been modified to produce novel and versatile plasmids pREP1 and pREP41. pREP1 contains a promoter derived from the gene nmt1. pREP41 contains a moderate-activity promoter (Pnmt41), whereas pREP81 contains a weaker promoter (Pnmt81). pREP vectors that contain ura4+ along with Pnmt1, Pnmt41, and Pnmt81 are named pREP2, pREP42, and pREP82, respectively [57]. The dominant selection marker genes kan, nat, hph, and bsd, which confer resistance against the specific antibiotics G418, clonNAT, hygromycin B, and blasticidin S, respectively, are used routinely during chromosomal integration [66, 67, 68, 69].
Other important kinds of vectors of S. pombe are those of the pRI series generated from vector pREP, which were produced by deleting the ars1 origin of replication sequence, and it has been used for the creation and expression of a single copy gene integrated into the chromosome [70].
The pYZ vectors are derivatives from the pREP series, which were designated for general purposes of cloning and large scale random gene cloning, as well as for allowing positive identification of cloning gene insertion and fusion to the GFP gene for analysis of gene expression. The pYZ vectors were constructed by inserting an E. coli α-peptide (position 239–684 on the pUC19 plasmid) of the lacZ (β-galactosidase) in opposite orientation to the Pnmt1 on the pREP series, leading to the complementation of the lacZΔM15 deletion in E. coli strains such as DH5α or JM105 [56, 71, 72].
The pREP1, pREP41, pREP81, and pSGA plasmids were generated from the pREP series called pYZ1N, pYZ41N, pYZ81N (N represents an additional Not I site), and pYZ3N-GFP, respectively. In those vectors, the distance between the Pnmt1 and the ATG start codon remains the same as in the pREP vectors, and the promoter strength is unchanged [71]. The pYZ vectors have been useful because they were designated to produce a correct positive identification of cloning gene, fusion to the GFP, and large-scale random gene screening. The versatility of the pYZ vectors has allowed their use in numerous researches. HIV-1 vpr is a virion-associated viral protein of about 12.7 kD, whose function is required for efficient viral infection of nondividing mammalian cells such as monocytes and macrophages [73].
The HIV-1 protease (PR) is a viral enzyme encoded by vpr gene that was initially expressed in S. pombe from pREP1N. Vpr makes proteolytic processing required to the production of viral enzymes and structural proteins and for maturation of infectious viral particles [74].With the aim to improve the functional studies, HIV-1 vpr gene was cloned in the pYZ vectors. The vpr gene was fused to GFP in the pYZ3N-GFP vector and expressed in the yeast, where Vpr localizes to the nucleus of fission yeast cells. Expression of the vpr gene from the pYZ1N vector allows the analysis of the effects on cell morphology, the cell cycle G2 arrest, and cell killing [75].
In the molecular analysis of the Zika virus (ZIKV) infection, a large-scale molecular cloning and functional characterization of the viral proteins were performed. The Zika virus (ZIKV) is the causal agent of the microcephaly and the Guillain-Barré syndrome after the viral infection. However, there is insufficient knowledge about how ZIKV viral proteins are involved in cell damage. So, S. pombe was used to identify ZIKV factors responsible for the ZIKV-mediated cytopathic effects as well as the pathogenic factors associated with the viral infection. By cloning the 14 coding-genes into the pYZ3N including the N-terminal GFP, it was possible to determine the subcellular localizations (nuclear, ER, Golgi, and cytoplasm) of ZIKV proteins expressed in a wild-type fission yeast strain, SP223 [70]. Importantly, seven ZIKV proteins affect cellular proliferation, which would be related to the microcephaly. So, ZIKV-induced microcephaly was proposed due to the intrauterine growth restriction, reduced cell proliferation, reduced neuronal cell layer volume, or cell death/apoptosis. Also, it was observed that prM, C, M, E, and NS4A proteins cause cell-cycle dysregulation because of cell cycle G2/M phase accumulation. These findings allow to follow the study of ZIKV infection.
Other interesting series of vectors are those that were produced as the pREP-X vectors that lack an ATG start codon [76]. Between them, pREP3X (promoter strength high), pREP41X (promoter strength medium), and pREP81X (promoter strength low), the three vectors lack tags and used Leu2 as marker. The pSLF vectors contain N-terminal or C-terminal triple hemagglutinin (3× HA) epitope tag. Between them, pSLF173 (promoter strength high), pSLF273 (medium), and pSLF373 (low), all of them contain 3xHA as tag and use ura4+ as the selective marker and the inducible promoter nmt1. From the pREP-X series were constructed several vectors with the purpose of being utilized for high-throughput functional analysis of heterologous genes in S. pombe such as pDS vectors that add GST taggings [50] as well as pSGA vector that includes GFP fusions.
There are many expression vectors constructed containing a destination cassette suitable for high-throughput cloning of target genes via the gateway system. There are vectors with N-terminal tagging such as the pDES173N, 273 N, and 373 N series, which add a 3XHA tag with the ura4+ gene as marker, and the vectors were constructed from the pSLF173, 273, and 373 vectors. The pDES175N, 275 N, and 375 N series add a GFP tag with the LEU2 marker, and those plasmids were built from the pSLF175, 275, and 375 vectors. The pDES177N, 277 N, and 377 N vectors add a GFP tag using ura4+ as marker selection. The pDES5XN, 45XN, and 85XN series add a RFP tag, with the LEU2 marker, which were derived from the pSLF5X, 45X, and 85X vectors. The pDES179, 279, and 379 series add a RFP tag, with the ura4+ marker, which were derived from the pSLF179, 279, and 379 vectors [77].
There are vectors with C-terminal tagging; those in the pDes173C, 273C, and 373C series add a 3XHA tag with ura4+ as marker, and the plasmids were constructed from the pSLF173, 273, and 373 vectors. The pDEs175C, 275C, and 375C series add a GFP tag with the LEU2 as marker, and those were constructed from the pSLF175, 275, and 375 vectors. The pDEs179C, 279C, and 379C series that add an RFP tag with the ura4+ marker were constructed from the pSLF179, 279, and 379 plasmids [77, 78]. These vectors exposed above lead the protein expression with N-terminal or C-terminal tagged, useful for the affinity purification or the functional analysis of target genes [77].
In 2013, an interesting series of vectors was described to PCR-based epitope tagging and gene disruption. The vectors developed were pFA6a-LEU2MX6, pFA6a-his3MX6, and pFA6a-ura4MX6. All of them were designed from the pFA6a-MX6-based plasmid (which contains antibiotic-resistance markers as kan) for amplification of gene-targeting DNA cassettes and integration into specific genetic loci, allowing expression of proteins fused to 12 tandem copies of the Pk (V5) (epitope from the P and V proteins of the paramyxovirus SV5), or 5 tandem copies of the FLAG epitope with a glycine linker. All vectors can use the LEU2, his3+, and ura4 + genes as selection markers. Also, some vectors as pFA6a-G9–5FLAG-kanMX6 and pFA6a-G11–5FLAG-kanMX6 were created, which were generated for studies of proteins when the direct epitope tagging compromises protein conformation and/or function. Other vectors were constructed to add a green fluorescent protein (GFP(S65 T)) or a monomeric red fluorescent protein (mRFP) genomic tagging as FA6A-GFP-bleMX6 [79].
Between the PK-tagging vectors are the pFA6a-6 × GLY-V5-(marker) and C-terminal FLAG-tagging vectors using KanMX6 and hphMX4 as markers. The FLAG-tagging vectors with N-terminal and C-terminal tags included the pFA6a-6 × GLY-FLAG-(maker), with kanMX6, hphMX6, natMX6, bleMX6, and his3MX6 as possible markers. Between the GFP-tagging vectors are pFA6a-GFP(S65 T)-(maker) and N-terminal and C-terminal GFP(S65 T)-tagging, which include kanMX6, hphMX6, natMX6, bleMX6, and ura4MX6. Also, some disruption plasmids as pFA6a-(maker), which has been used for gene deletions using kanMX6, hphMX6, natMX6, bleMX6, ura4MX6, his3MX6, and LEU2MX6, were constructed [79].
A novel system to cloning several DNA fragments, into a plasmid, is the Golden Gate shuffling method. Golden Gate cloning [80, 81, 82] is a modular cloning system and was set up for simultaneous overexpression of multiple genes. Some of the applications of the Golden Gate that have been tested in Pichia pastoris are the development of strain engineering, pathway expression, and protein production [83].
The use of this methodology for the construction of pREP1-type plasmids that expressed GOI-FPtag was reported S. pombe. To apply the Golden Gate cloning, several modules including promoters, tags, marker genes, terminators, and the gene of interest (GOI), which are cloned separately, are produced separately. They are digested with the enzyme BsaI that recognizes a specific sequence GGTCTC and cleaves any four-base sequence after it (such as nNNNN, mMMMM, and kKKKK) at 37°C but generates cohesive ends for various sequences. The Golden Gate method connects all the modules in the order desired in a single reaction. The cleaved fragments are joined by DNA ligase at 16°C. Once complementary four-base overhangs are connected, the site can no longer be cleaved with BsaI. The temperature shift is repeated up to 50 times until circular plasmids are efficiently produced. The system allows the assembly of up to eight expression units on one plasmid with the ability to use different characterized promoters and terminators for each expression unit [84].
In first place, modules were prepared using the pREP1 vector [70]. A segment from pREP1, which includes ars1 and Amp, was amplified by PCR with a pair of oligonucleotides containing BsaI and NotI sites. A typical expression plasmid for S. pombe is composed of six modules in total. The modules are a promoter, a terminator, a GOI, an FPtag fused at the N- or C-terminus, a selection marker such as an antibiotic resistance gene, and auxotrophic marker gene required to select colonies that harbor the expression plasmid. With this method, several plasmids were generated. The first plasmid was named pBMod-exv (colEI ori, Amp, ars1, NotI, and KanR sites), and this plasmid was the backbone of all vectors. Plasmids named pRGG (from pRGG-1 to pRGG-5) are expression vectors designed to express GFP-Atb2 from pREP-type multicopy plasmids. For the construction of pRGG-1, LEU2 was chosen as a marker module, whereas for pRGG-2, kan was chosen. To further demonstrate the convenience of the Golden Gate method, a series of plasmids of variable promoter strength were designed to express GFP-Atb2. The genetic elements included were the promoter (nmt1–41-81 and adh1–41-81 y urg1), an FPTag-N (GFP+ linker, mCherry+ linker, and CFP+ linker), an FPTag-C (linker+ GFP, linker+ mCherry, and linker+ CFP), GOI, and Terminator + marker (Tadh + Kan, Tadh + hpd, Tadh+nat, and Tadh+bsd) [84].
Recently, pheromone-inducible expression vectors for were developed S. pombe. By replacing the native Pnmt1+, the promoter regions of the sxa2+ and rep1+ genes were utilized to couple pheromone signaling to the expression of reporter genes for quantitative assessment of the cellular response to mating pheromones. The rep1+ and sxa2+ genes were chosen considering that sxa2+ mRNA increases more than 1600-fold upon pheromone perception in M-type cells [85, 86]. The EGFP open reading frame was placed downstream of the pheromone-inducible promoters, yielding pJR1-rep1-EGFP and pJR1-sxa2-EGFP, respectively [87].
In some cases of the heterologous protein expression, the better way to obtain the right protein production host is through its ability to secrete high titers of properly folded post-translationally processed and active recombinant proteins into the culture media. Proteins secreted in their native hosts will also be secreted in the culture medium. Some signal sequences used to secrete the protein into the extracellular space include α-MF and SUC2 invertase. Both are derived from S. cerevisiae α. α-MF is composed of a pre- and proregion and has proven to be most effective in directing protein through the secretory pathway. Other signal peptides to sorting are PHO1 P.p. acid phosphatase, SUC2 S.c. invertase, PHA-E phytohemagglutinin, KILM1 Kl toxin, pGKLpGKL killer protein, CLY and CLY-L8 C-lysozyme and syn., leucin-rich peptide, and K28 pre-pro-toxin K28 virus toxin, to produce molecules such as human interferon, α-amylase, α-1-antitrypsin, and human lysozyme [88].
One of the major problems to the correct production and purification of heterologous proteins from fission yeast is the proteolytic degradation of the recombinant gene product by host-specific proteases. To avoid that problem, a protease-deficient disruptant was constructed set by disruption of 52 S. pombe protease genes using the PCR-mediated single gene-targeted gene disruption method. This technique was used to delete the full open reading frame (ORF) sequence of each target protease gene, using ura4+ as the selection marker [89].
In the first place, the protease-deficient disruptant was obtained, which was amplified from genomic DNA of the S. pombe ARC010 strain, using appropriate adapter designed to fuse with the 5′ and 3′ termini of ura4 (1762 bp), respectively. Then, by fusion extension PCR, ura4 was sandwiched with the resultant PCR products to obtain the gene disruption fragment (2.2–2.3 bp). The resultant DNA fragments were then introduced into competent cells of the ARC010 strain, using the lithium acetate-based transformation method. Then, the efficient protecting activity of protease of the mutant strains was analyzed. A chromosome-integrative hGH expression vector using the pXL4 plasmid was constructed [89].
To analysed the levels of the secretory production of human growth hormone (hGH), that its known to be a proteolytically sensitive model protein. The results indicated that some of the resultant disruptants were effective in reducing hGH degradation. Although in some cases, added inhibitors of proteasas like Antipain, bestatin, Chymostatin, E-64, Leupeptin, pepstatin, Phosphoramidon, EDTA, aprotininto avoid protein degradation were necessary. Eight protease coding genes useful for reducing degradation of recombinant proteins [isp6 (subtylase type 9 proteinase), pgp1 (endopeptidase), psp3 (subtylase type peptidase), sxa2 (serine carboxypeptidase), ppp51 (aminopeptidase), ppp53 were identified (zinc metallopeptidase), ppp60 (metalloprotease) and ppp80 (peptidase)], the use of a strain lacking the aforementioned enzymes allowed a high level of recombinant hGH production. This publication raised the need to evaluate different proteases to identify those that are the best candidates for the production of recombinant proteins, as well as for functional screening, specification, and modification of proteases in S. pombe [89].
In relation to the methods for the transformation of S. pombe, the lithium acetate and polyethylene glycol-based transformation of plasmid DNA are the most popular and temperature stresses. With these methods, it is possible to achieve transformation efficiencies between 1.0 × 103 and 1.0 × 104 transformants per microgram of the plasmid with 108 S. pombe cells [90, 91].
The use of mutants to analyze the function of genes has been a tool widely used in S. pombe. In this yeast, several types of mutants have been produced such as the temperature-sensitive mutants with conditional defects in the ability to participate in some cellular process in the cell cycle, cytokinesis, lipid metabolism, or DMSO-sensitive [92]. The use of temperature changes to impose a restrictive condition is a strategy widely employed. But, there are methods such as altered sensitivity to drugs, pheromones, and changes in ionic strength, among others. For mutational analysis, the haploid state offers the advantage to observe the effect of specific mutations [93].
In the case of the essential genes, a lethal phenotype is frequently observed. To achieve the study of essential genes, there are two strategies. First, the mutations or gene deletions are created in the diploid state and then the synthetic lethality is studied in the haploid state. Sometimes, it’s possible to observe a slow-growth phenotype, in which haploid cells can partially survive without function of the inactivated gene. Second, the creation of the conditional lethal mutations allows to study a relatively normal gene function under permissive conditions, and then the loss of function is observed under nonpermissive conditions. The most used conditional mutants are the temperature sensitivity, sensitivity to DNA-damaging agents, sensitivity to drugs and inhibitors, and dependence on amino acids or certain carbon sources for viability. Three methods highly used to produce mutants are gene knockouts, random mutagenesis, and site-directed mutagenesis [94].
The CRISPR/Cas system is a bacterial defense mechanism, and its main function is to identify and degrade exogenous nucleic acid sequences [95]. CRISPR-CAs is organized in an operon, which codes the CAS proteins, and a series of identical repeated sequences separated by other sequences known as spacers, which are recognized by intruding DNA molecules [96]. A part of the nucleic acid stranger is incorporated into the spacer’s zone of the operon using the Cas proteins, which degrade the strange DNA. Next, the transcription of CRISPR-Cas generates a precursor CRISPR-RNA or pre-crRNA, which is then processed to generate crRNAs of small size, which are complementary to the sequence of the foreign DNA. In the last known phase of interference, Cas proteins, using as a guide to crRNAs, detect intruding sequences and degrade them [96].
The CRISPR/Cas technology allows to identify a specific segment of DNA, remove, or replace it using always the same tools: a duplex RNA with the copy of the DNA to be identified (sgARN) and a short sequence adjacent to the proto-spacer (PAM) that will bind to DNA and stabilize the protein Cas9, protein with endonuclease activity, and helicase guided by the sgARN that separates and cuts the two strands of DNA. A Cas9-gRNA plasmid expressing the active Cas9 enzyme and sgRNA, as well as another plasmid with donor DNA for each deletion are required. The CRISPR-Cas technology allows targeting of multiple genetic manipulations to the same strain, it avoids indirect physiological effects, and it limits the perturbation of the local chromatin and transcriptional environment to the gene manipulation of interest. In fission yeast, this technique has allowed to produce genetic modifications as point mutation knock-in, endogenous N-terminal tagging, and genomic sequence deletion [97].
Recently, a web-tool called CRISPR4P CRISPR for Pombe or CRISPR Pombe PCR Primer Program was developed as freely available from the website (
A gap-repair-based CRISPR/Cas9 procedure allows to efficiently knockin a point mutation in fission yeast. The rpl42-P56Q mutation confers cycloheximide resistance (CYHR) [100]. Employing this technique, a CCC codon for proline was changed, and with the use of a pair of 90-nt complementary oligos as donor DNA, the gap repair procedure achieved a high editing efficiency (84%).
Using the CRISPR-Cas9, yeast strains, functional and successfully complemented with the markers ura4-D18, leu1-Δ0, his3- Δ0, and lys9-Δ0, were created. To achieve the goal, all the components were assembled with the “BsaI-pad,” a single 42 bp region containing two BsaI cutting sites to produce the plasmids pYZ182, pYZ183, and pYZ184 with nmt1, nmt41, and nmt81 cassettes, respectively. Using that design, the marker genes ura4, leu1, his3, and lys9 were integrated separately. Later, the plasmids were transformed into yeast [101].
Recently, the type VI CRISPR system, Cas13a from Leptotrichia shahii (LshCas13a), was employed to introduce genetic changes on the DNA, disrupting or editing to target and knockdown endogenous gene transcripts with different efficiencies in S. pombe [102].
RNA interference (RNAi) is a highly conserved eukaryotic gene regulatory mechanism, which uses small noncoding RNAs to mediate posttranscriptional gene silencing as a host defense mechanism. It was described that S. pombe has the entire RNAi machinery (Dcr1, DICER ribonuclease; the Rdp1, RNA-dependent RNA polymerase 1; and the Ago1, Argonaute family member). In S. pombe, the role of the RNAi pathway on the heterochromatin assembly has been widely studied [103]. RNAi plays a role in regulating expression of Tf2 retrotransposons, and it is also involved in the RNAi-dependent heterochromatin assembly by the Hsps, Hsp90 and Mas5 (a nucleocytoplasmic type-I Hsp40 protein).
siRNA is generated by the Dicer family endoribonuclease Dcr1, from double-stranded noncoding RNA that is complementary to heterochromatin. The siRNA duplex is loaded onto a non–chromatin-associated complex called Argonaute, small interfering RNA chaperone (ARC), which contains the Ago1 endoribonuclease. The loading of the siRNA duplex onto the Ago1 subunit requires the two ARC-specific subunits, Arb1 and Arb2, which also inhibit the release of the passenger strand [104]. Thus, this complex changes its subunits’ composition to form a chromatin-associated effector complex called RNA-induced transcriptional silencing (RITS) [105]. The RITS complex is composed of Ago1, now binding single-stranded siRNA as a guide for target recognition, and the two RITS-specific subunits: Chp1 and Tas3. Chp1 uses a chromodomain to recognize H3K9me, whereas Tas3 bridges Ago1 and Chp1 [106].
To analyze the role of the RNAi in fission yeast, the lacZ fission yeast system was employed. With this system, it was possible to know that the gene inhibition is dependent on the dose of the antisense RNA, the size of the antisense transcript, as well as the targeted region. Any of them can affect the efficacy of target gene inhibition. The generation of dsRNA through either intermolecular or intramolecular hybridization is central to make the antisense RNA-mediated gene silencing in S. pombe [107]. As a genetic tool to analyze the function of genes, the ura4-based RNAi-based selective assay was developed using a repressible thiamine promoter [108]. The RNAi must be optimized in order to know the minimum requirements to achieve the knockdown of a specific gene. U-HP construct was produced as a hairpin complementary to 200 bp of ura+ gene expressed from the nmt1 promoter and integrated at ars1 on chromosome 1. U-HP silences ura4+ inserted nearby to centromere 1, but not the endogenous ura4+ gene. Interestingly, in S. pombe, exogenous siRNAs can only silence efficiently in trans, when the target locus is near endogenous sites of heterochromatin.
An interesting proposal to analyze the role of the siRNAs in S. pombe was achieved with the development of a GFP-HP construct. This system was generated under control of the Pnmt1, and it contains two GFP open reading frames arranged in an inverted orientation, around the first intron from the rad9 gene. When it was probed, it was demonstrated that GFP-HP induces trans-silencing of target genes. GFP siRNAs generated by the expression of a GFP-HP can act in trans to establish heterochromatin on target genes bearing homology to GFP siRNAs and silencing their expression. This silencing does not require other manipulations, such as deletion of eri1+ or increased expression of Swi6HP1, a heterochromatin component, to promote RNAi-mediated silencing in trans [109].
The yeast two-hybrid system (Y2H) is a method widely employed to study the physical interaction of proteins by the downstream activation of a reporter gene. Considering that many eukaryotic transcription factors are organized in a modular way with at least two domains, it is possible to separate them into their domains [110].
In this assay, two plasmids are created; the first is named the bait plasmid including the DNA-binding domain of a transcription factor joined to one of the proteins to analyze and it is named Bait. In this vector, a selection marker is included such as HIS3, ADE2 (Gal4 system), or LEU2 (LexA system with binding sites for the DNA-binding domain). The second vector is named prey including the activation domain of the transcription factor joined to the second protein to study in the interaction, named Prey. As in the other vector, a different selection marker is included. When the Bait and Prey proteins are put together by protein interaction, they restored the organization of the transcription factor, and then they can activate the transcription of the reporter gene as the E. coli lacZ gene. The transcription factors more frequently used are Escherichia coli LexA protein and the yeast Gal4 protein, as well as herpes simplex virus VP16 protein and the B42 acid blob from E. coli [111].
Gal4 is a transcriptional activator in yeast that binds to UAS (upstream activation domain), a specific DNA sequence, and activates transcription in the presence of galactose. The separation of Gal4 in two fragments produces N-terminal DNA-binding domain (DBD) and C-terminal transcriptional activation domain (AD), but did not activate transcription in the presence of galactose until both domains are associated to reconstitute a fully functional Gal4. Some disadvantages of the assay consider that in some cases, it’s necessary to modify the bait proteins because a protein with both DNA-binding and transcriptional activating properties is possible to be found. Some fused proteins may not be able to enter or be expressed in the yeast nucleus. The GAL4 BD has its own nuclear localization signal (NLS). If the GAL4-based Y2H system fails, the interaction could be analyzed and detected successfully using a LexA-based Y2H system [110, 111].
The Y2H system has been widely used. In S. pombe, its use in the searching of the new determinants of aging was reported. Chen et al. described a method to select long-lived mutants from S. pombe bar code-tagged insertion mutant library (each insertion had a unique sequence tag called a bar code produced by random barcode). With this strategy, it was possible to identify an insertion mutation or deletion in the cyclin gene clg1+ that extended the chronological aging of the yeast. At the same time, it was determined that depletion of Clg1p also decreases the cyclin-dependent kinase Pef1p and an extended longevity was observed. To analyze if the phenotype was produced by direct or indirect contact, a yeast two-hybrid analysis and immunoprecipitation assay were performed [112].
To the assay, the entire pef1+ ORF was fused to the Gal4p DNA-binding domain and the entire clg1+ ORF was fused to the Gal4p activation domain. A physical interaction was observed between Clg1 and Pef1. To perform this assay, the pGBT9-Pef1 and pGAD424-Clg1(full length) or pGAD424-Clg1(1–590) plasmids were constructed and transformed into the Saccharomyces cerevisiae two hybrid indicator strain Y187 (MATα, ura3–52, his3–200, ade2–101, trp1–901, leu2–3112, gal4Δ, met-, gal80Δ, MEL1, and URA3::GAL1UAS-GAL1TATAlacZ, Clontech). Positive transformants were selected on complete medium plates without leucine and tryptophan at 30°C for 3 days. The reporter gene lacZ expression was probed from five individual colonies from each transformation and was patched on plates that require both plasmids for growth and incubated at 30°C for 2 days. Then, the coimmunoprecipitation was performed with FLAG-tagged Clg1p, which was expressed in cells that also expressed triple HA (3HA)-tagged Pef1p [113]. Using Western blotting of FLAG-Clg1p immunoprecipitates revealed the presence Pef1p-3HA. Chen et al. concluded that Clg1p interacts with the cyclin-dependent kinase Pef1p in S. pombe cells. In addition, a third Pef1p cyclin named Psl1p was identified. Genetic and coimmunoprecipitation assays indicated Pef1p controls lifespan by downstreaming the protein kinase Cek1p [114].
DNA microarray is an orderly set of segments of genes that are immobilized on a surface called chip. The DNA arrangements allow the massive study of the gene expression of an organism, and it allows to know the differences of gene expression between two samples of RNA in a given cellular condition. In cells that present some mutation or elimination in some genes or cells derived from individuals with some infectious disease or not, the microarrays allow the identification of sets of genes related to the gene or genes under study or the condition of disease. Comparing RNA prepared from diseased cells and normal cells can lead to the identification of sets of genes that play key roles in diseases. Genes that are overexpressed or underexpressed in the diseased cells often present excellent targets for therapeutic drugs.
The application of DNA microarray technology requires a genomic library conformed by a set of DNA segment derived from each of the genes of the model of interest, which is generated from PCR products or synthetic oligonucleotides, as well as the design and construction of the arrangement, to determine the physical location and accurate identification for the analysis and interpretation of gene expression data. Microarray analysis requires total RNA extraction from control and the problem obtained by any strategy optimized for certain cell type [115]. Total RNA control and the problem should be submitted to retrotranscription incorporating uracil marked with a fluorescent molecule as dUTP-Cy3, dUTP-Cy5, dUTP-Alexa 555, dUTP-Alexa 647, and biotin, among others. The labeling of the cDNA must be differentiable between the two tissues to be analyzed [116]. The hybridization of the microarray containing probe sets that represent a finite number of transcripts is carried out. Fluorescence reading is obtained with a microarray reader. The quantification of the signal produced by the fluorescence of the spots allows to calculate for each point the mean density value of the nucleotides marked cDNA (g. e. of Alexa555, Alexa647) and the average value of the background. To identify the genes expressed differentially in the experiment, it is necessary to perform a statistical analysis, from the normalization of the data. The goal is to analyze those genes that move away from normalization through the value of Z [117]. The genes with the value of Z > 2 present a statistically significant change between the experimental condition and the control (genes with greater or lesser expression). [116]. Easy and useful software for data analysis of microarrays is GenArise (computer unit of the Institute of Cellular Physiology of UNAM (
From the data that record a significant change, it is necessary to determine its association to some biological processes by clustering analysis for gene expression [118].
With this molecular tool, it was possible to analyze in fission yeast the effect of Spc1, a mitogen-activated protein kinase in the stress responses. Spc1 is an activator of transcription factors that control gene expression in response to extracellular stimuli and is also known to interact with the translation machinery. Using microarrays of Affymetrix GeneChip Yeast Genome 2.0 Array, it was possible to know the set of genes that is regulated by SPC1, and this analysis was carried out without and with a stress condition to evaluate the effect of the wild-type SPC1 kinase and Spc1K49R, a mutant of this enzyme. Spc1 and Spc1K49R were separately overexpressed in S. pombe cells, and gene expression was compared with the control cells (which are transformed with the empty with the Pnmt1). Interestingly, only 42 genes were found with differential expression after Spc1 overexpression, while 132 genes were found to be differentially expressed after Spc1K49R overexpression. Some of the genes up-regulated after Spc1 overexpression were Mitogen-activated protein kinase sty1 and M cell-type agglutination protein mam3. The downregulated genes were NAD-dependent malic enzyme, meiotic cohesin complex subunit Rec8, and aph1 bis(5’-nucleosidyl)-tetraphosphatase. Between genes differentially expressed after Spc1K49R overexpression, those upregulated included pheromone p-factor receptor, RNA-binding protein involved in meiosis Mei2, MAP kinase Spk1, cell agglutination protein Mam3, M-factor precursor Mfm1, and M-factor precursor Mfm3. And some downregulated were serine/threonine protein kinase Gsk3, RNA-binding protein Sap49, and Argininosuccinate lyase [119].
In 2016, the role of the putative NO dioxygenase SPAC869.02c (Yhb1) and the S-nitrosoglutathione reductase Fmd2 was analyzed. Both proteins are NO-detoxification enzymes. In the study, it was found that exogenous NO protects S. pombe cells against H2O2-induced oxidative stress by inhibition of Fe(3+) to Fe(2+) conversion, upregulation of the H2O2-detoxifying enzymes, as well as downregulation of the MRC genes. Transcriptomic analysis was carried out with an Affymetrix Gene Chip Yeast Genome 2.0 Array [120].
The fission yeast S. pombe generally reproduces by mitosis. To know the role of the fhl1 protein in meiosis, a microarray analysis of the fhl1∆ strain was performed. Interestingly, it was found that nitrogen starvation-response genes are controlled by fhl1. Some of them are genes of mating and sporulation such as isp4, mfm1, mfm2, Mat-Mc, ste4, ste11, map1, map3, mei2, and mcp7 [121].
Next-generation sequencing (NGS) involves the parallel mass sequencing of thousands of DNA fragments. Sample processing for NGS can be summarized as follows: First, nucleic acid extraction (DNA or RNA). Second, selection of the type of NGS sequencing (targeted sequencing, whole exome sequencing, and whole genome sequencing). Third, library generation by DNA fragmentation, ligation of adaptors, and amplification and sample enrichment. Fourth, template generation or cluster generation according to the platform of sequencing. Fifth, sequencing (using a specific platform as Illumina, PacBio). Sixth, data analysis. Data analysis includes the quality evaluation of the sequence, alignment to reference sequence to identify some possible variations such as single nucleotide polymorphism (SNP) or insertion-deletion (indel) identification, phylogenetic or metagenomic analysis, as well as the identification, interpretation, and classification of pathogenic variants [122, 123].
Splicing is an essential step in eukaryotic gene expression. Introns are excised by the spliceosome, composed of five uridine-rich small nuclear RNAs (U1, U2, U4, U5, and U6 snRNAs) and several polypeptides. To characterize the U2·U5·U6 complex of S. pombe, cell lysates were obtained. A large-scale isolation of the U2·U5·U6 complex was performed using double-affinity purification using a split TAP-tag approach [124], with protein A attached to U2 snRNP protein Lea1 (U2 A′ in humans) and calmodulin-binding peptide (CBP) attached to U5 snRNP protein Snu114 (U5 116K in humans). After the purification of the complexes, the content of protein and RNA associated to the U2·U5·U6 complexes was analyzed. By denaturing PAGE and high-throughput sequencing (RNAseq), the presence of U4, U1, and heterogeneous higher molecular weight species was shown. In addition, the U2·U5·U6 snRNA complex contains excised introns, indicating that it is primarily the ILS (intron lariat spliceosome) complexes. The protein content of the ILS complex of S. pombe was similar to the spliced product of humans and the ILS complexes assembled on single pre-mRNAs in vitro from Saccharomyces cerevisiae [112].
There are some other techniques to study several aspects of the physiology of S. pombe. Chromosome conformation capture (Hi-C) is a technique widely used to identify long-range chromatin interactions. The spatial organization of mitotic chromosomes with the greatest compaction during mitosis is an interesting aspect of the cell cycle. In S. pombe, it is known that condensin, a structural maintenance of chromosomes (SMC) family member, has a role on the chromatin architecture. Biochemical studies have been applied to discover the more relevant points of the mechanism. By chromosome conformation capture (Hi-C), it was demonstrated that condensin is able to replace short-range local contacts in the interphase with longer-range interactions in the mitosis. Condensin achieves this by setting up longer-range, intrachromosomal DNA interactions, which compact and individualize chromosomes. Even local chromatin contacts are constrained by condensin during mitosis [125].
Finally, it is necessary to mention that Rallis & Bähler offered to the world pombe community an excellent review showing the relevance of S. pombe in the eukaryotic studies employing a wide genome screen and phenomic assays, ranging from growing conditions to metabolomics [126, 127].
Schizosaccharomyces pombe is an excellent model to study highly conserved processes between eukaryotes, its versatility, ease of manipulation, its accessibility to genetic manipulations, making it a great model system increasingly used by a growing scientific community interested in fission yeast. At the same time, this interest has promoted the technological development, the implementation, and the continuous improvement of new molecular tools that when applied to S. pombe will allow to elucidate new mechanisms of cellular processes with potential application to the Eukaryotic kingdom including the human being.
Children’s experiences with digital technologies actually involve an increasing quote of young users (also defined as “digital natives”) who are born and are developing in environments in which new digital technologies are widely available [1]. This currently occurs from early infancy, due to the rapid diffusion of touchscreen devices among younger children (or “touch generation”; [2, 3]). Children aged 2–4 years actually are able to use touchscreen devices, such as tablets or smartphones, to play or watch movies, and often parents themselves introduce kids to use them in boring social situations (i.e., in the pediatrician’s waiting rooms or in the restaurant; [4]). On the basis of the most recent report on worldwide diffusion of the Internet among young people [1], one in three users is estimated to be a child or teenager (under 18). Generally children use digital technologies in their home, particularly younger children, with intense and prolonged activities especially on weekends. Children often use their digital technologies at school at least a day a week (almost 30% among 9–11 years), although it is prohibited in many countries by school regulations. The access to digital technologies is expanding among young generations, even if many inequalities of resources remain between developed or developing countries [1]: for example, it has been estimated that in Africa (Ghana) children mainly use 0.9 mobile devices to connect to the Internet, against 2.9 in South America (Chile) or 2.6 in Europe (Italy). Similarly, only 12% of children in Africa (Ghana), 21% in the Philippines, and 26% in Albania can connect to the Internet at school, against 63–54% of children in other South America or European countries, such as Argentina, Uruguay, or Bulgaria. This reality raises several questions on how to guarantee the young generations the opportunities offered by new technologies (for studying, enhancing skills, socializing, etc.), protecting them from potential dangers of digitalized world (i.e., contacts with unknown people, exposure to violent/pornographic contents, etc.). In fact, although children grow in a reality permeated by new media, they are not automatically “digitally literate,” that is, able to juggle the digital world and to reflect on it. Studies show that not only young users, but also teenager users “have difficulties in finding, managing and evaluating information, managing their privacy online and ensuring their online personal safety […]and may thus vary in their digital skills” ([5], p. 186).
Together with their children, parents themselves are largely exposed to media experiences in many fields of their life. Digital technologies have quickly changed the way in which family members communicate, enjoy themselves, acquire information, and solve daily problems. Parents are also the first mediators of children’s experiences with digital tools: they have the task of integrating their use into ordinary routines (play, entertainment, learning, mealtime, etc.), promoting constructive and safety uses. Digital parenting describes parental efforts and practices for comprehending, supporting, and regulating children’s activities in digital environments. A growing research on digital parenting identified the main approaches that can allow parents to “mediate” children’s activities with digital technologies [6, 7, 8]. According to Vygotsky’s theory of child development and his concept of proximal development zone [9], parental mediation can be considered a key aspect in facilitating the interactions between children and new media. The proximal development zone is an intermediate area between what the child is able to do alone and what he/she can learn thanks to the guidance of others. In the course of a shared activity, the support and the help are adapted so that the child can improve his/her skills and gradually assume responsibility for acting alone. However, the activities that take place in the virtual environments of the web, unlike the experiences in the real environments, can reverse the relationship between the competent person (the adult) and the learner (the child). Today’s children have an early, almost “intuitive” approach to digital technologies, so in some cases they can become active agents towards their parents. When children’s knowledge and digital competence (e.g., functions/benefits of a new app) overcome that of parents, many shared experiences can be child-initiated, and children can also perform some forms of support and digital teaching to parents. This reverse socialization [10] seems to be a peculiar feature of digital experiences, and it poses new challenges to parental role. Reverse socialization describes all situations where children possess a better understanding or more advanced skills than adults. This gap between generations is more marked in low-income families or low-educated parents who possess limited resources and access to digital technologies [11]. However, over the past years, many parents have developed adequate knowledge and technical skills to share digital experiences with their children [3, 12]; they appreciate benefits of the web and strive to comprehend its complexity.
A common difficulty that parents actually encounter derives from the diffusion of “portable” devices (smartphone and tablet) that children start to use in early infancy (under the age of 2; [13]). Later, due to unlimited Wi-Fi access and enhanced connectivity, children insert activities with mobile devices into many daily routines, for example, during mealtime, school homework, conversations with parents, or before sleeping [14]. Particularly, parents worry about the “pervasiveness” (or ubiquitous) of mobile technologies in daily activities [15], and they fear that an effective guidance and control over them may decrease. Studies with large samples of young digital users (9–16 years old) in many European countries have compared parents’ opinions before (2010 Eu Kids Online Survey; [12]) and after (Net Children Go Mobile; [3]) the diffusion of mobile devices. After 4 years, many parents declare that they know less about their children’s online activities and have more difficulties to closely monitor children’s usage (e.g., time spent connected). Interestingly, parents now are more aware of the risks of using the web [16], and they prefer to talk to children about Internet security (e.g., do not leave personal data online or block unknown people) rather than limiting or prohibiting Internet use [17]. Parents can encourage or limit the use of digital technologies to children according to the opportunities or danger they attribute to them. Since parents themselves are regular, sometimes enthusiastic, users of digital media, their digital skills and confidence and daily frequency of usage (or overuse; [18]), together with beliefs about digital world [3], are all crucial factors that researchers have begun to explore systematically.
Each parent has beliefs, that is, convictions and personal opinions, regarding the usage of media by children, such as their usefulness or damage, or the age at which children should use them. Beliefs are the cognitive dimension of attitudes, guiding individual’s behavior and choices. When parents raise their children, they act and make choices for them following their own perceptions of what is desirable or what they positively value for their child’s development [19]. Although parents are not always aware of their beliefs, these influence parent-child interaction and the child’s opportunity to learn, do experiences [20], and develop digital skills [5]. Parental beliefs are important aspects of parenting and family microsystem, together with factors such as parent’s history and education, socioeconomic status, and culture.
Parents possess personal ideas about modern technologies: they can be considered a source of entertainment/relaxation or a learning tool [21, 22]; conversely, for other people, PC, tablet, and smartphone can be harmful to children’s health (such as sleep problems, obesity, etc.; [23]), for social risks (such as contacts with unfamiliar or social isolation; [24]), or because they interfere with parent-child activities and time spent together [25].
A qualitative study [26] shows that parents have more pessimistic (70.55%) than optimistic opinions (29.45%) on the Internet use by primary school children: for example, parents worry about the excessive time spent online, the interference in face-to-face conversation, or that children lack of skills and maturity in dealing with some contents suitable for older children (such as violence, sex, or drug-related contents). Other worries concern negative consequences on learning and academic performance (i.e., reduced attention span), physical development (i.e., prolonged sedentary activities), social skills and peer interactions (i.e., fewer opportunities to “learn to play together”), and child’s well-being (i.e., using smartphone to overcome boredom). Interestingly, many parents fear losing control over their children’s online behaviors. Conversely, the positive beliefs concern positive effects of digital technologies on child’s entertainment, communication and learning, access to information, and enhancing of child’s skills (such as brain functioning, self-regulation, autonomy, critical attitude, etc.).
Other researchers [27] explored parent’s perceptions about positive (i.e., they are shared by generations) or negative impact (i.e., they expose family privacy to risks) of social media—such as Facebook or WhatsApp—on family open communication. Teenagers are intensely involved in social media use, but adults also are regular users. On the one hand, parents use social networks to communicate; on the other hand, they fear that they negatively impact family relationships, for example, through the phubbing phenomenon (i.e., ignoring someone or interrupting a conversation or mealtime to check the smartphone). Authors found that parents’ perceptions are a meditational variable between the collective family efficacy (i.e., the perceived efficacy to manage family relationships, to support each other, etc.) and the openness of communication: “it is not only the actual impact of social media on family systems that matters but also parents’ perceptions about it and how much they feel able to manage their children’s social media use without damaging their family relationships” (p. 1).
Parental beliefs may influence the degree to which parents give opportunities or restrict their children’s media use, but beliefs should not be considered the “cause” of behavior towards children. Researches show that parents’ positive beliefs (e.g., “the tablet improves reading skills”) are associated with favorable attitudes, co-using approach, communication, or suggestions to enhance their child’s appropriate use of the Internet [28]. For example, when parents think that smartphones are useful tools (i.e., they promote child’s intelligence and knowledge), they more often allow their preschool children to use them (i.e., at the restaurant), and children become regular users, spending more time (at least 2 h a day) with smartphone activities [29]. Conversely, parents who attribute negative effects to digital media tend to limit activities to children (i.e., put time limits or react for smartphone overuse); in turn, these restrictive behaviors can influence how much the children use these devices [28]. Therefore, the influences of parental beliefs on child’s behaviors are not directed, but they are mediated by parental practices and other factors such as parental education or involvement with mobile device (“attachment”; see, e.g., [30]) that can intervene.
Parental beliefs include also self-efficacy [31, 32], that is, parent’s sense of competence in their own digital skills and in managing their children’s technology usage. An example of parental self-referent estimation of competence is “I won’t bother setting parental controls or passwords because my kids will “hack” around them” (cfr. [33]). In many studies, parental self-efficacy is positively associated with active parental practices: when parents feel confident about their Internet skills, they more often are involved in or monitor their children’s media activities [6]. Recently Shin [34] distinguishes general self-efficacy (the confidence to be a good parent; [35]) from two self-efficacy domains assessing parental beliefs more strictly related to digital tasks: parental “media competency” in using media technology (such as sending/receiving email with a smartphone) and “perceived control over mediation strategies” (the degree to which the parent feels to be able to guide or modify their children’s behaviors on smartphone). All these domains of parenting self-efficacy are associated with each other [34], suggesting that perceived competence on their own digital skills can positively influence parents’ involvement with children (e.g., discussing about smartphone use).
Sanders et al. [33] found that when parents are confident to have adequate digital skills, they more often intervene (i.e., with rules and reinforcement strategies) with their children. Parental self-efficacy also influences parental opinions about technologies and how they talk about them with children [33]. Moreover, parental perception of influence in managing technologies decreased with preadolescents that generally are seen as more self-regulated and reluctant to the parental control than younger children. These findings suggest the importance to recognize the influence of child characteristics (such as age, technology usage, perceived competence, etc.) on digital parenting.
Initially studies on parental engagement in children’s activities with media assumed as theoretical basis the traditional parenting styles [36, 37]. According to Darling and Steinberg [38], parenting styles are defined as the context (or emotive climate) in which parents raise and socialize their children, and they are distinct from practices, that is, the distinct actions contingent to the child’s behavior (e.g., scolding when the child uses the smartphone during mealtime). As it is well known, two main dimensions of the parent’s behaviors, and their natural variations along a continuum, describe the styles: responsiveness/warmth (involvement, acceptance, and affect that the parent expresses towards the child’s needs) and demandingness/control (rules, control, and maturity expectations for the child’s socialization). Parenting styles derive from the combination of these variable dimensions: authoritative parenting (high warmth and high control, e.g., parents listen to the child’s wishes, but they put clear limits to the child’s behaviors); laissez-faire parenting (low warmth and low control; the parents are detached from the needs expressed by the child; they did not give rules or limits to child’s behavior); authoritarian parenting (low warmth and high control; parents expect the child to obey; they neither discuss nor listen to the child’s opinions and can react with harsh discipline); and permissive parenting (high warmth and low control; parents are very affectionate, but they lack in guidance through rules and give few limits to the child’s behavior).
Studies that applied these “classic” parenting styles to children’s behaviors with new communication media did not provide convincing results [39]. As an alternative to the “broad” parenting styles, a description of specific media-related practices is more useful in empirical studies for exploring the link between parental behaviors and child outcomes (e.g., time spent online). Therefore, researchers strove to identify the key dimensions of parental warmth/control more strictly referred to children’s behaviors on the Internet or new media (Table 1). These Internet parenting styles are more strictly linked to children’s actual use of digital technologies, for example, low parental control predicted more time of Internet usage by school-aged children [8].
Style dimensions | Item (examples) |
---|---|
Parental control | Supervision: “I’m around when my child surfs on the Internet” |
Stopping internet usage: “I stop my child when he/she visits a less suitable website” | |
Internet usage rules: “I limit the time my child is allowed in the Internet (e.g., only 1 h a day)” | |
Parental warmth | Communication: “I talk with my child about the dangers related to the Internet (costs, addiction to games, computer viruses, privacy violation, etc.)” |
Support: “I show my child “child friendly” websites (library, songs, crafts, school website, etc.)” |
Dimensions of the internet parenting style (adapted from [8], p. 89).
Parenting style dimensions seem influenced by parents’ individual characteristics such as gender, instruction, beliefs, or prior experiences with digital technologies. For example, in Valcke et al. [8] study, mothers are more controlling but also warmer than fathers, both dimensions associated with an authoritative style. In other studies, younger fathers and those who use the Internet more frequently with their teenagers are higher in control [40]. Parental instruction and experiences with digital technologies are other important variables: higher educated parents are more involved and high in control, probably because higher instructional levels also correspond to greater parents’ competence with the Internet [8].
The first studies explored parenting styles related to Internet usage at home, but more recently other authors explored the influence of digital parenting styles on children’s usage of mobile devices (tablet and smartphone). Konok et al. [30] found that children (3–7 years old) who use the devices for more time every day have parents who are more permissive (e.g., they talk with children about applications on devices, but have low levels of demandingness), more authoritative (e.g., they give time limits, but they do not block the use because they expect the child to regulate himself), and less authoritarian (i.e., the parent restricts and prohibits mobile use). Interestingly, these parenting styles are also associated with parental beliefs about positive/negative consequences of early media usage: parents who have higher permissive or authoritative digital style declared more beneficial (i.e., skill improvement, entertainment, and early learning of digital skills) than negative effects (i.e., reduced time for other activities, developmental problems, and danger/addiction) for children’s mobile usage.
Digital parenting styles change also according to children’s characteristics, such as age [41], self-esteem [42], emotion regulation [43], or behavioral problems [44] that can intervene, mediating the link between parenting and children’s actual behavior with digital technologies. Particularly, styles vary and accommodate with children’s age: authoritative parents during infancy become more permissive with older children [41]. Overall, these findings reappraise the idea that there is a linear, cause-effect relationship between parenting and child outcomes on digital behaviors, but bidirectional and transactional parent-child influences [45] should be considered.
Alternatively to digital parenting styles, many researchers adopted parental mediation as perspective for exploring parental influences on children’s digital behaviors. Parental mediation refers to “the diverse practices through which parents try to manage and regulate their children’s experiences with the media” ([7], p. 7). Parental mediation strategies were initially introduced in empirical studies as a potential factor influencing children’s use of television [46] and videogames [47]. These studies, exploring how parents can effectively reduce excessive exposure or enhance children’s self-regulated behaviors, inspired the following researches on digital technologies. Actually in literature two broad mediation approaches are distinct: enabling (or instructive) mediation and restrictive mediation [16]. These strategies are only partially similar to those parents who adopt “traditional” media: for example, co-viewing is a mediation strategy generally applied to television use [48], but it is difficult to apply it to portable media (particularly, smartphone and tablet) that children often use alone or outside the home environment. As a consequence, parents can feel worried because they cannot effectively control their children’s media use and involvement in digital life [11, 49].
The (a) enabling mediation is also defined as “active” or “instructive mediation” in that parents engage different activities with the aim to enhance their child’s appropriate use of the digital technologies: for example, they explain to him/her how to use a media device, talk about the contents of new app/websites, or play a videogame together (co-use mediation). Nevertheless, in many empirical studies, (b) co-use (or co-viewing mediation) does not imply parent-child conversations, but the parent is present when the child displays the activity with the media without discussing the content [13]. The (c) restrictive mediation is characterized by a strict attention to rules and control to the child’s digital activities: for example, parents decide when the child can have his/her tablet, pose time restrictions, or react when the child uses the smartphone too long. The (d) technical restriction is a particular kind of restrictive approach adopting software applications or other technical tools to control the child’s activities (e.g., installing filters on PC for children’s safety). Nevertheless, parents rarely use them and declare they prefer child-directed strategies, such as giving explanations or sharing the device [6].
Active mediation is the most frequent approach adopted in European families with 9–16 years old children, whereas restrictive mediation strategies are more common with younger children [16]. Interestingly, when children are interviewed about the mediation approach adopted in the family, they agree with their parents’ responses [12].
All mediation strategies are linked with changes in children’s digital behaviors, for example, less time exposure with online activities [12], or reduction of negative outcomes (i.e., aggressive behaviors, overuse, etc.; see [50]), but their efficacy is relative and it changes as a function of the child’s development (i.e., age and digital skills) and his/her actual activity with media. Active mediation is linked with positive outcomes (such as social and cognitive skills), particularly with younger children (0–3 ages): for example, during video/movie watching, parents stimulate attention, comment, or pose questions to children, giving them occasions for language exposure and cognitive and digital learning [51]. Nevertheless, we cannot link children’s outcomes uniquely to a distinct mediation strategy, since parent-child interactions are complex and many contextual or individual factors can intervene. Parents often use a combination of mediation strategies, and they change the mediation approach according to the activity the child is doing (e.g., using the tablet for school homework or for visiting Facebook; [11]).
Other authors explored the influence of family sociocultural factors. For mediation to be effective to guide children’s experiences in the web, parents need to have themselves knowledge and skills of the new digital media (see Section 4 in this chapter). Particularly in conditions of sociocultural disadvantage, parents may lack basic digital skills [52], or they may not be able to explain to children how digital reality works and rapidly changes [53]. Unlike the traditional media (such as television or video game console), parents can give a difficult task to assure a help or guide children with the ever-changing technologies. Recently, Nikken and Opree [11] found that mostly low-educated, low-income, and single parents are likely to experience low competence and greater insecurity with new devices (such as electronic screen), declaring that it is difficult to apply co-use or active mediation strategies with their young children (1–9 ages). In addition, Warren and Aloia [49] found that when parents perceive high stress levels, the restrictive mediation and the discussions with children about contents and the use of media increase.
Parental mediation strategies may change according to their child’s age and his/her digital skills, but longitudinal studies are scarce in literature. Developmental changes have been observed from childhood to adolescence: active mediation strategies more often are adopted with younger children, whereas restrictive mediation fades with older and adolescents [17]. Parents generally expect greater autonomy and self-regulation skills from adolescents, and the influence of some parental strategies decrease over time: for example, the efficacy of restrictive strategies (i.e., rules for time or negative consequences for overuse) in reducing screen time decreases with older children [33]. From a developmental perspective, particularly the effects of restrictive approach are unclear. Some studies evidence that restrictive strategies (such as limiting access to media) are effective with younger children [6], but not with older kids. Adolescents can perceive parental control/limitations as a violation of their needs (i.e., self-determination, privacy, peer relationships, etc.) and react with increased online activities [54].
After all, parents wish their children can develop self-regulation, critical view, and awareness of opportunities or risks of digital technologies. In many studies, parental active mediation—for example, discussing with children issues such as cyberbullying, sexting, and online frauds—is more effective than restrictive mediation in reducing risks [16, 55]. Conversely, the efficacy of restrictive mediation must be considered relatively, since in literature both positive and negative associations with online risks emerge [56]. Mascheroni et al. [57] comment, “While restrictive mediation can be effective in reducing children’s exposure to online risks, it has numerous side-effects, because it limits children’s opportunities to develop digital literacy and build resilience and discourages children’s agency within the child-parent relationship. Enabling mediation, instead, encompasses a set of mediation practices (including co-use, active mediation of internet safety, monitoring and technical restrictions such as parental controls) that are aimed at empowering children and supporting their active engagement with online media. The question is, then, how to ensure children’s access to online opportunities while protecting them from potential harmful effects.”
Interestingly, parents adopt their approach according to their child’s competence in digital technology use (digital literacy). In line with a bidirectional model of parent-child influences [45], not only parenting influences child’s behaviors, but also the child’s actual behavior or perceived digital competence influences parental behaviors. Generally, restrictive mediation strategies are more often adopted with less digitally skilled children, but this approach could be counterproductive: limiting online activities for protecting the child from risks, in turn, can deprive him/her to opportunities for developing adequate digital skills [5]. Conversely, parents more often use active mediation strategies (e.g., they share experiences or talk about media) with skilled children than with children who have scarce competencies [58].
The predominance of online activities in the life of many children often worries parents, who observe that spending much time online removes children from face-to-face relationships and social activities. Empirical studies confirm the negative effects of Internet unsuitable use on social participation, since high levels of online activities are associated with few friends, reduced offline relationships [59], and increased loneliness [60]. Particularly loneliness, that is, social isolation and lack of intimacy with close friends, was found to be strongly associated with Internet excessive use [61]. However, causal relationship between Internet excessive use and loneliness is still under investigation [62], in an attempt to understand if loneliness can be the antecedent or the consequence of the individual’s excessive involvement with Internet activities. Two alternative hypotheses have been proposed to explain the link between poor social involvement, feeling lonely, and the development of problematic Internet use in children. According to the first hypothesis, loneliness is one of the main antecedents of excessive online activities, together with low self-esteem, poor social skills, social anxiety, and frequent conflict with parents. Some authors (e.g., [63]) hypothesized that adolescents who feel lonely or experience high anxiety in face-to-face social situations may use social networks and online exchanges more frequently than non-lonely adolescents. According to this “compensation hypothesis,” they are increasingly involved in Internet activities that provide alternative experiences for social life. The second hypothesis assumes that time spent online causes loneliness and social withdrawal, isolating and depriving people of real social experiences. Therefore, loneliness can be considered as a possible outcome of Internet overuse [64], like when prolonged activities online reduce time spent with family and friends. Finally, there are studies that did not confirm the link between loneliness and Internet problematic use [65] or that evidence some positive consequences on individual socioemotional well-being. For example, contradicting the assumption that using the web impoverishes social life and increases isolation, in some studies higher levels of Internet activities are positively associated with social connection and perceived support. Unfortunately studies with children and adolescents are still lacking, but the attention among researchers is growing [60, 66].
Given the paucity of research with adolescents, we conducted an unpublished study1 to explore the relationships among excessive Internet use, preferred online activities, and adolescent’s perceived loneliness. In addition, we hypothesized that among adolescents better parent-child communication and higher parental emotional availability were positively related with less time spent online and less frequent online activities. In fact, studies indicate that parent-child communication and parental involvement play a protective role to excessive online activities [67]. A community sample of 177 high school students (66% females), aged 16–22 years old (M = 18, DS = 1.01), completed a questionnaire measuring the sense of loneliness (UCLA Loneliness Scale; [68]) and the Compulsive Internet Use2 Scale (CIUS, [69]) for assessing problematic involvement in Internet activities. Daily frequency of favorite online activities (chatting, e-mailing, visiting social networking sites, listening to music, watching videos, playing online games, etc.) was also measured. Regarding parenting factors, adolescents filled out (a) the Lum Emotional Availability of Parents questionnaire (LEAP; [71]) assessing adolescent’s perception of parental responsiveness, sensitivity, and emotional involvement and (b) two scales (derived from [70]) measuring the frequency of communication (how often the adolescent communicates with parents about his/her online activities) and the quality of parent-child communication (the adolescent feels understood, or comforted, or taking seriously from parents when he/she talks about Internet activities). In our study loneliness was not associated with Internet compulsive use (CIUS scores), but with specific online activities. Adolescents with higher loneliness levels reported higher frequency of music listening, but they declared less access to social networks (such as Facebook). This result contradicts the hypothesis of social compensation assuming that the teenagers use online exchanges to replace the sense of loneliness in real life [61]. An alternative explanation, proposed by others [72] is that a process downward with a “spiral pattern” is activated: loneliness leads to a decrease in social involvement which in turn increases the sense of isolation. Interestingly, those who spent more time online and were problematic users (higher CIUS scores) were more frequently involved in solitary activities, such as watching videos, listening to music, playing games offline, and visiting social networking sites. Perceived emotional availability from the father (but not from the mother) was negatively related with time that adolescents spent online. Teenagers who perceived greater emotional availability from both parents used the Internet more often for working on school projects and homework or doing search. A better quality of communication with parents is associated with less use of the Internet for gambling and online games. Overall these results confirm a virtuous relationship between quality of family communication, emotional availability of parents, and productive use of the web.
An interesting evidence emerging from empirical literature is the protective role of parent-child communication for preventing Internet unsuitable use in children [73]. Conversely, Internet excessive use is associated with low quality of communication in the family [74]. Particularly with teenagers, the open and effective parent-child communication is a key dimension of family relationships and climate. Assuming a bidirectional perspective of adolescent-child influences, some authors focus on the role of youths’ self-disclosure and spontaneous communication on parenting. Stattin and Kerr [75] claim that parental efforts to monitor adolescent’s activities or to discuss about them are ineffective if teenagers do not trust their parents and if they are not willing to open up spontaneously. Parental monitoring on children’s activities can be less effective when it is parent-driven (e.g., the parent tries to follow the child’s activities on Facebook) than when it is child-driven, that is, activated by children’s self-disclosure and open communication. Conversely, when parents try to control teenagers’ online communication (e.g., the friends on Facebook, the photos posted on Instagram, etc.), parent-child conflicts increase, and adolescents can perceive parental behaviors as an obstacle to their autonomy or an intrusion to privacy [76].
Van den Eijnden et al. [70] identify two key dimensions of parent-child communication about children’s digital behaviors. The first parenting practice refers to the frequency of communication about Internet usage (e.g., “How often do you and your parents talk about who you have Internet contact with?”), whereas the quality of communication about Internet use measures adolescent’s perception of mutual respect and acceptance during conversation (“When my parents and I talk about my Internet use, I feel taken seriously”). Authors explore how these parental behaviors, together with other Internet-specific parental practices (rules about time online, rules about contents, reactions to excessive use), link to compulsive Internet use (CIU) in adolescents. Findings from their longitudinal study are particularly interesting, showing a protective effect of the quality of communication, but not of frequency of communication, on the risk of developing CIU. In other words, a good quality of parent-child communication about the use of Internet decreased the risk of CIU (6 months later), whereas this relationship was not observed for the frequency of parent-child exchanges about adolescent’s online activities. Authors discuss these findings by highlighting the bidirectional nature of parent-child influences. When adolescents show compulsive Internet behaviors, the frequency of parent-child communication decreases. Probably gradually parents get discouraged and give up the idea of achieving a positive change in their child’s problematic behaviors through frequent conversations.
Regarding the parental rules about online activities, studies evidence some mixed results. When parents give their children rules about the content of the Internet, the compulsive use of web decreases; conversely, strict rules about time allowed for online activities seem to be counterproductive, linking to compulsive Internet behaviors in children [70]. Moreover, considering the child’s influences on parent’s behaviors, it is possible that when the child remains connected online without time limits, her/his behavior in turn stimulates stricter rules by parents. Other studies evidence that parental rules about Internet use are less influential on their children’s behaviors than their parents’ behaviors. Liu et al. [77] found that when parental behaviors are consistent with parental rules regarding digital technologies and the Internet (e.g., the smartphone must not be used during mealtime, personal data cannot be given online, etc.), the rules negatively predict Internet problematic use in adolescents. This result reminds us the importance of educational consistency (i.e., rule-behavior agreement) from parents. Conversely, when parental rules and parental behaviors do not agree, only the parents’ behaviors are positively predictive of children’s excessive Internet use. According to social learning theory [78], a parental modeling process intervenes, that is, an observational learning in which the parent’s behavior acts as antecedent for similar behavior in the child. Therefore, parents act as a role model for their children’s digital behaviors, and young children learn how and under what circumstances to use a mobile, for example, the smartphone, observing parents’ activities with that device. Interestingly, studies show that the time parents spend with computers positively relates with time spent by their children [79]. Similarly, parental involvement in favorite Internet activities (visiting social networking sites, video streaming, etc.) is positively associated with the same activities engaged by children. In addition, as some researchers remind us “it is not only overt parental behavior (i.e., digital device use) but also attitudes and emotions that can be modelled for children to imitate” ([30], p. 4). Taken together, these findings suggest that parents’ agreement and modeling of adequate behaviors are crucial factors for promoting self-regulation and safety use of digital technologies in young children.
Today’s reality is widely digitized, and it offers people of all ages opportunities for socialization, amusement, learning, job, and knowledge that were unthinkable until a few decades ago. Precisely in the weeks in which the authors were engaged in the revision of this chapter, COVID-19 pandemic was involving more than 130 countries in the world. The lockdown and restrictions at home quickly changed daily activities of children and parents, transferring to the screen of the devices many activities previously carried outdoor (school lessons, play with peers, etc.). It is still too early to know what impact the epidemic will have on children’s physical and mental health, but the attention of professionals and researchers is not lacking [80]. Surely during COVID-19 screen time has increased exponentially in the families: in some ways for the parents it was a relief, because through the Internet children continued their school courses and contact with peers. In addition, children avoided boredom through videogames or website dedicated to music, creativity, etc. On the other hand, the intensive online activities have renewed parents’ concerns about the well-known risks [23, 81], such as increased sedentary and physical inactivity, prolonged use at night, sleep disorders, isolation, and escape in digital world by teenagers.
Following social distancing and the temporary closure of schools for limiting COVID-19 infection, the Ministries of Education in many developed countries quickly activated online courses and other websites for distance learning. These online solutions have the aim to guarantee children’s right of instruction but also to mitigate the negative effects of home confinement [82]. However, online courses shift the teaching from school to home and make the parents a resource for support and effective learning. The question is: what can be the role of parental mediation and digital competence? As the authors know, there are no empirical studies on this topic, but previous studies with primary school children showed negative associations between parental control, interference in homework, and children’s learning [83]. Currently, in many cases teachers expect parents to ensure that their children connect on time and follow the video lessons, so parental support could be useful, but tensions and parent-child conflicts can also occur. There is also the risk that parents may help children, interfering with digital learning or impeding them from carrying out the assigned activities independently. Close attention and research effort are needed for comprehending how this aspect of digital parenting works, supporting parents in their efforts and ensuring a good home learning to children.
In line with the available studies before COVID-19 [4], we believe that during lockdown the digital activities satisfy children’s basic psychological needs, such as socialization and emotional support by the family (grandparents and cousins) and other significant people (teachers and peers). Social media facilitate the expression of emotions (such as fear and sadness), self-disclosure, and the keeping of romantic relationships by adolescents particularly [84]. Video calling and regular contacts through smartphone have been recommended as an important source of reassurance in the cases of isolation of the caregiver or family due to prevention of COVID-19 infection or recovery [85].
What probably becomes necessary in the time of COVID-19 is a renegotiation of family routines, that is, a balance between screen time and other moments of family life. In this regard, the WHO [85] recommends that parents maintain regular routines for children (school/learning, free time/relaxing, bedtime, etc.) and also to create new opportunities for joint activities (such as co-use for creative, amusing, or physical activity in front of the screen). With young children, many shared activities offer also a context to express and communicate their feelings (both fears and wishes) in a supportive parental relationship. Even in actual COVID-19 circumstances, we believe that parental behaviors (such as self-limiting screen time for smart working, chatting, or gaming) are more influential than restrictive mediation or limitations imposed to children.
Having the digital knowledge and the skills to move in the digital world, without suffering the dangers, is not a matter of age, but of education and learning, that is, digital literacy. It is a serious responsibility towards the new generations and a complex challenge for which the adults (parents, teachers, psychologists, or educators) do not feel prepared. As Martin ([86], p. 135) reminds us: “Digital literacy is the awareness, attitude and ability of individuals to appropriately use digital tools and facilities to identify, access, manage, integrate, evaluate, analyze and synthesize digital resources, construct new knowledge, create media expressions, and communicate with others, in the context of specific life situations, in order to enable constructive social action; and to reflect upon this process.” Currently, parents’ difficulties stem from the fact that they—as digital users—have different levels of involvement, technical skills, and beliefs that influence mediation practices towards their children. If parents feel less skilled or worry about unknown dangers of the web, they could activate more restrictive practices, but rarely they will be able to critically discuss with their children in a constructive manner. In addition, parents believe not to be up to their children in juggling in the digital world, in pursuing technological innovations, or in protecting children from danger or media abuse. Sometimes parents consult the websites for suggestions on how to effectively manage kids in their digital activities, but information disseminated through the websites is not always scientifically founded (fake news). The researcher Danah Boyd [87], in describing the complexity (“It’s complicated”) of teenagers’ life on the web, claims that the media magnify the virtues (the “superpowers”) of digital natives, but at the same time they trigger parental fears talking about serious dangers such as Internet addiction, sexual enticement, or incitement to suicide. Conversely, rarely parents turn to professionals for advice. A study [28] conducted with families of very young children (under 7 years) shows that parents choose the type of help (professionals such as pediatricians, or friends and family) based on the child’s problems and his/her digital activities. The professionals are consulted if the child is an only son or he/she uses the media too long. Parental sense of competence in managing the child’s activities increases if parents are confident of the usefulness of the media (e.g., educational games for learning) and if there are more kids in the family. Parents turn to friends and family for advice when they have a negative view of the effects of the media. This result makes us reflect, but unfortunately there are not many similar studies.
A correct parental mediation of children’s digital activity must build on the information and recommendations that come from the scientific community. The American Academy of Pediatrics [2] has taken a clear stance for prudent and moderate use of the web in infancy (0–5 years) and has prohibited touchscreen device use under the age of 2. The careful use of these devices at such an early age is crucial for the infants’ brain and social development. However, in contrast to these professional recommendations, often parents themselves introduce babies to media use during infancy (e.g., to “take calm” the kid, or to stop whims and cry; [30]). Young children spent daily an amount of time with screen media (iPod, smartphone, video game player, etc.) that grows during infancy (42 min under 2 years and 2 h/39 min at 2–4 years, respectively; [88]). The risks for excessive screen exposure are extensively confirmed in literature and particularly the negative consequences for early users who may present physical problems (such as obesity), developmental difficulties (i.e., language or learning), and unhealthy routines (low sleep quality) (Figure 1).
Developmental risks associated with excessive media exposure (from [88]).
The recommendations for effective parental mediation on children’s digital activities are unequivocal [2]: (a) avoid the use of digital devices before 18–24 months with the exception of video chatting in the presence of the parent; (b) do not allow the child (18–24 months older) to use the devices alone and for more than 1 h a day; (c) do not press for an early use, the child will spontaneously approach the media when ready; (d) help the child apply what he/she learns from using the device to the real world; (e) know that in infancy, direct experiences, manipulation, and unstructured play are crucial for the child’s brain and for social, cognitive, and linguistic development; (f) void the vision of fast programs, with too many distracting elements, or violent contents that the child is unable to understand; (g) avoid using devices to calm the baby, an hour before bedtime; and (h) constantly monitor the media contents to which the child is exposed. Finally, the experts (pediatricians and psychologists) turn also to the industry that produces media devices, so that it adopts a scientifically founded and more ethical approach, for example, installing apps (such as connection stop or automatic shutdown during night hours) that can protect very young children from the risks of overuse.
Therefore, parent education interventions are necessary both to disseminate scientific knowledge on the influence of new technologies on children’s health and development and to help parents to cope with the challenges of digital reality. Parent education cannot be reduced to merely correcting ineffective parenting practices or to a list of instructions on what the parent should do. In fact, all studies indicate that the effectiveness of mediation strategies (restrictive or active approach) is relative, because parental practices interact with the characteristics of both adults (digital skills, beliefs, and activities on the media) and children (age, development, digital literacy skills, etc.). Instead, professionals should help parents to improve and adjust their guidance according to children’s age and developing skills. This is possible to be realized if parents also increase their knowledge and digital skills (media literacy programs), given the importance of these factors in parenting. Less skilled parents, or those who fear the unknown pitfalls of the web, are more likely to intervene only on restricting or prohibiting children’s activities. Conversely, “it is likely that more skilled children and parents are more free to explore and benefit from online opportunities, while also building up resilience against harm by meeting a degree of online risk” ([16], p. 19).
Digital parenting is a very complex and “complicated” task not only because the digital technologies rapidly change, but also because they offer children multiple experiences (learning, communication, socialization, entertainment, etc.) that influence their development, but which are not entirely overlapping to the experiences that take place in the real environment [89]. Particularly, digital natives have the opportunity to know the reality and themselves, developing their own identity [76], with a multiplicity of means and without the supervision of the traditional agents of socialization, primarily the parents (or the teachers). With the awareness of how difficult it is to give definitive answers about the advantages or dangers of digital technologies, more effort is needed from researchers. More evidence-based studies are needed, to understand how technological progress is changing the psychological (neurocognitive, emotional, and social) development of young digital users. However, despite the growing diffusion of digital tools in infancy, studies with very young children are still lacking. Particularly, future research could benefit from longitudinal studies to which to explore the relationships between parenting and children’s experiences in digital environments, their opportunities, or risks.
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