Released this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\\n\\n
We wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
IntechOpen is proud to announce that 179 of our authors have made the Clarivate™ Highly Cited Researchers List for 2020, ranking them among the top 1% most-cited.
\n\n
Throughout the years, the list has named a total of 252 IntechOpen authors as Highly Cited. Of those researchers, 69 have been featured on the list multiple times.
\n\n\n\n
Released this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\n\n
We wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
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1. Introduction
Hemp plastic is a bio-based composite that may vary in composition and applications. The components of plant material that provide valuable properties and usually constitute 25–65% of the composite [1] are bast fibers. These cells are developed within hemp stem and have specific morphological and mechanical parameters. Hemp fibers, same as fibers of flax, ramie, nettle, and some other fiber crops, are distinguished by high-cellulose content [1, 2, 3]. The quality of such fibers varies depending on both the processing and the properties developed in planta [4, 5, 6]. We will consider the biological determinants of hemp fiber quality, comparing primary and secondary hemp fibers. Being developed within the same plant, primary and secondary hemp fibers differ significantly in morphology, cell wall organization, and as a consequence in quality [7, 8, 9], giving the possibility to understand how developmental processes influence quality parameters.
2. Origin of primary and secondary fibers in the hemp stem
The usually considered primary and secondary hemp fibers are bast fibers, meaning that they belong to phloem and are located closer to stem periphery than cambium (Figure 1). Primary phloem fibers located behind the epidermis and collenchyma are larger than the secondary ones (Figure 1A, B). Secondary fibers located closer to the cambium and arranged in compact bundles, the number of which grows during plant development (Figure 1A, C, D). Hemp stems, same as in many other dicotyledonous plants, have also fibers located within xylem, but they do not form bundles, are mixed with other cell types (vessels, parenchyma) and have different properties and applications [10]. Xylary fibers are not considered further in this chapter.
Figure 1.
Primary and secondary phloem fibers within hemp stem. Cross-section of hemp stem, stained with toluidine blue. Stem bottom of plant at (A) flower formation and (D) seed maturation stages; (B) primary and (C) secondary phloem fiber bundles. Bar scale = 100 μm (A and D), 20 μm (B and C). C, cambium; Co, collenchyma; E, epidermis; P, parenchyma; Pf, primary phloem fibers; Ph, phloem; Sf, secondary phloem fibers; and X, xylem.
By definition, primary fibers originate from primary meristem: procambium, in the region close to apical meristem [8]. Their initiation is coupled to leaf trace formation and occurs within the very top millimeters of the developing stem (Figure 2A, B). Apical meristem provides the increase of stem height [11]. The more downwards the stem, the older the primary fibers. Thus, developmental gradient of primary fiber can be observed along the same stem. However, it is better to characterize it, analyzing fibers at the same height through plant development, same as it was done for flax [12]. This is because primary fibers located at different stem heights originated at different periods of plant development and might be differentially affected by various endogenous and environmental factors. For example, primary fibers from the stem bottom are known to be shorter than the ones from the middle stem part [13]. Fibers traced at the same stem height through plant development have a similar background and can be correctly compared. The total amount of primary fibers in the hemp stem calculated from the total volume of primary fiber bundles and average volume of a fiber cell is around 700–800 thousand [14].
Figure 2.
Scheme of initiation and intrusive elongation of primary and secondary phloem fibers in hemp stem. (A) The top of hemp stem (2 mm) [8] with primary fibers at the stages of initiation, symplastic growth, and beginning of intrusive elongation. (B) Primary and secondary phloem fibers at different stages of development, their characteristics, and localization within the stem (on the left); initiation of primary and secondary fibers from procambium and cambium, correspondingly, and morphology of fibers at different developmental stages (on the right); the length of fibers at different stages is scaled to real proportions, while the diameter/length ratio is larger than its actual value. (C) Changes in the number of primary and secondary phloem fibers on transverse section in the developing hemp stem.
Secondary fibers originate from secondary meristem: cambium that largely provides cells for stem thickening [11]. Secondary fibers start to develop more than half a meter away from stem top (Figure 2B), as demonstrated for several monoecious varieties [7, 8, 14, 15, 16]. Their developmental gradient can be traced within the stem radius: the closer to the cambium, the younger the secondary fibers. Within hemp stem, secondary fibers may form several distinct concentric layers separated by other cell types. The number of such rings may reach 3–4 (Figure 1D). The total amount of secondary fibers in the hemp stem is around 2 million, much higher than the number of primary fibers [14]. From that, only small proportion of cambium cells (around 10%) give rise to secondary phloem fibers [8].
3. Intrusive elongation of primary and secondary fibers
Plant fibers are distinguished by their extreme length [11, 17, 18, 19]. It is mainly attained by the special type of cell elongation: intrusive growth [11, 17, 19]. This process has an enormous, though often overlooked, effect on fiber yield and quality. Intrusive elongation is characterized by the higher rate of a cell growth as compared to its neighbors. It is distinct from symplastic (also called coordinated) growth [20], when all involved cells increase their surface with the same rate, as it happens in the growth zones with most of the tissues [11]. Fibers are the classical example of cells performing intrusive elongation [11, 17, 21, 22]. During such growth, a fiber has to split middle lamellae of the cells on the way and intrude between them. Herewith, new contacts are made along the increased fiber surface, so that the stem tissues do not fall apart.
Primary phloem fibers of hemp stem start intrusive elongation rather soon after being initiated from meristematic cells [8] (Figure 2A, B). Only shortly primary fibers grow symplastically with the surrounding cells and then increase the rate of elongation. By symplastic elongation, primary phloem fibers of hemp attain the length of 200 μm [8] (Figure 2B). The start of the primary fiber intrusive growth is marked by the formation of the so-called “knee”: the flat tip of the symplastically growing cell is transformed into the tapered one to effectively intrude the surrounding tissues [8, 19, 22]. In hemp stem, such structures are observed at 1.8–2.0 mm from the very top (Figure 2A, B). By means of intrusive elongation, primary phloem fibers of hemp stem increase their length roughly hundredfold so that their average length gets around 18,000 μm. The volume of a fiber is increased even more, since intrusive elongation is accompanied by the increase of fiber width, so that the cell diameter gets 30 μm instead of 3–4 μm at the end of symplastic growth [8] (Figure 2A). The final fiber dimensions of primary phloem fibers in hemp may differ depending on the variety and growth conditions. The final length of such fiber is reported between 5000 and 100,000 μm, with modal class being around 20,000 μm, and variations in fiber diameters, which could vary between 15 and 40 μm [2, 15, 23, 24, 25]. Thus, the ratio of cell length and cell width in a primary fiber of hemp may constitute several thousands, being among the highest in plant cells.
The duration of intrusive elongation can be traced by the increase in the number of fibers on the stem cross-section. As described above, the formation of new primary phloem fibers occurs only at the region close to apical meristem [8], then during intrusive elongation, the number of primary fibers in the cross-section of hemp stem increases till approximately 60–90 mm from the apex (Figure 2C); end of the intrusive elongation occurs within this stem region. Further downward the stem, fibers do not elongate and their number on cross-section does not increase with time.
Elongation of fibers goes bidirectionally, as indicated by the presence of “knees” at both cell ends [8, 26]. The intrusive elongation of primary fibers goes for several days and is completely split in time with cell wall thickening, which starts later in the course of fiber development [8, 12, 26]. After the start of cell wall thickening, intrusive elongation cannot be restored any more.
During intrusive elongation, primary fibers initiated at different leaf traces reach each other, this leads to the formation of fiber bundles (Figure 3). The structure of the fiber bundles is almost exclusively determined by fiber intrusive growth [8, 12, 22]. The longer the fibers in the course of elongation, the thicker the fiber bundles.
Figure 3.
Formation of primary phloem fiber bundle within the hemp stem. Fibers formed within one leaf trace are given in one color; fibers formed in different tree leaf traces are given in different colors (black, yellow and red); fibers formed within other leaf traces are given in white. The scheme illustrates the formation of primary phloem fiber bundle of hemp stem and participation of fibers formed within tree leaf traces in this process.
It is due to intrusive growth that fibers in a bundle are so tightly packed to each other and have no intercellular spaces. The large surface of the contacts between neighboring fibers formed during intrusive elongation permits them to stay together during the retting process [4, 23, 27]. These tight contacts may be further reinforced by the special type of the “glue” between cells—pectic compounds that are constituents of middle lamellae and primary cell walls [28], and may have peculiarities in composition in fibers performing intrusive growth [22]. In the course of intrusive growth that leads to the enormous increase of cell surface, pectins are actively deposited and modified. Low-molecular mass phenolic compounds may also be involved in strengthening of interactions between polymers of fiber primary cell wall [29].
Unfavorable environmental factors, like drought, may influence the extent of fiber intrusive elongation. This would lead to the shorter individual fibers and thinner fiber bundles in the region of the stem that contained elongating fibers [30]. For primary phloem fibers, this region is located at the top of the developing stem, usually constitutes 6–9 cm, and corresponds to the maximum length of individual fibers that are elongating in this region. After the end of unfavorable conditions, this stem portion would contain fewer fibers on the cross-section and would remain the weaker part of the whole bundle until the end of plant development.
Secondary fibers do not have the stage of coordinated growth, since the stem portion ceases elongation before they are initiated from the cambium. The final length of secondary fibers is achieved solely by intrusive elongation (Figure 2B). The final dimensions of secondary fibers are much lower than that of primary fibers. Their average length accounts for 2000–8000 μm, while the cell diameter is usually about 6–8 μm [8, 10, 15, 24]. However, these smaller dimensions are still considerably larger than those of cambium initials, the length of which in hemp stem was estimated to be around 250 μm [8]. As mentioned above, secondary phloem fibers start to emerge approximately at the middle part of hemp stem (600–700 mm from the apex) and their initiation continues down to stem bottom. The amount of secondary fibers on the stem cross-section increases toward the base of the stem due to both initiation of additional fibers from cambium and intrusive elongation of already existing fibers (Figure 2C). Due to smaller size of secondary fibers, their proportion in total yield of bast fibers in the hemp stem does not exceed 45% [10], despite the higher amount of individual cells [14]. This demonstrates the importance of the intrusive elongation stage for the final yield of bast fibers.
The bundle of the secondary fibers within the certain concentric ring is formed by a very similar process as described above for primary fibers (Figure 3). Fiber initiated from a cambium cell elongates and joins other intrusively growing fibers that originated from different cambium initials. The structure of fiber bundles is determined not only by the extent of fiber length increase, but also by the direction of elongation. The analysis of fiber-enriched peels demonstrates that the bundles of primary fibers in flax look like “straight columns,” while the bundles in hemp form the ramified net (Figure 4). The degree of ramification is especially high for secondary fibers [8, 31]. Such difference is due to the “joint” or “individual” behavior of elongating fibers. In flax, the elongating fibers follow the way made by the “oldest” fiber in the forming bundle. This can be due to the special mechanical properties of middle lamellae that have been already split. For primary fibers of hemp, the situation is rather similar, but some fibers escape from the bundle, elongating in a different direction and may reach another bundle, leading to formation of a ramified net. Such difference may be related to the large increase of stem circumference in hemp due to secondary growth, which is far less in flax.
Figure 4.
Comparison of phloem fiber bundles in flax and hemp. Structure of (A) primary phloem fiber bundles of flax, (B) primary and (C) secondary phloem fiber bundles of hemp [8] on the strips peeled off from the stems. Fiber bundles of hemp frequently split and merge along the stem forming numerous anastomoses (marked by arrows) unlike fiber bundles of flax. Bar = 100 μm.
The importance of intrusive growth for fiber yield and quality demands the approaches to regulate it. However, the mechanisms of intrusive growth are understood quite poorly. Nothing is known about the mechanisms that trigger and stop it. The peculiarities of fiber physiology at this stage of development are barely characterized. The reason for that is the difficulty to study this process, since it occurs within the depth of tissues and has never been reproduced in vitro. Fibers at this stage of development, being quite long cells, have only primary cell wall and can be easily damaged during sample preparation [26, 32]. This makes it quite difficult to obtain intrusively growing fibers for analysis by methods of biochemistry or molecular biology. The important step to identify molecular players involved in various stages of hemp fiber development was performed by the analysis of the transcriptome in hypocotyls of different age [33] and in different parts of young stems [34], both of which may contain intrusively growing fibers. However, the analyzed samples contained complex mixture of tissues and the early stages of fiber development were not fully identified. The indication of the stage-specific participants of fiber intrusive elongation in hemp may come from the analysis of whole transcriptome of intrusively growing fibers of flax. Such fibers were obtained by cryosectioning of stem and further, laser microdissection of fibers specifically at the stage of intrusive elongation [35]. However, elucidation of the mechanisms that perform and regulate the intrusive growth of fibers still has a long way to go.
To summarize the importance of fiber intrusive elongation, it (1) determines the final size of each individual fiber, (2) leads to the formation of fiber bundles and dictates their structure, (3) provides the tight contacts between the fibers that help to withstand retting process, and (4) provides the large surface for the further deposition of thick cell wall that is the major component of mature fibers.
4. Cell wall thickening
The mechanical properties of mature individual fiber largely come from thickened cell wall. In hemp fibers, cell wall may get 15 μm thick and occupy over 90% of cell cross-section [15]. The thick cell wall is not uniform and contains several layers with distinct properties. After the primary cell wall that is formed during fiber elongation, the secondary cell wall layer (S1) is deposited (Figure 5). The rest of the two layers of the thickened cell wall in hemp fibers are often also considered as secondary cell wall layers and named correspondingly, S2 and S3 [15, 31]. However, they differ significantly in composition and structure from S1. As revealed by fluorescent microscopy, LM10 and LM11—antibodies specific for xylan [36]—label only the outer layer of fiber cell wall [37]. Electron microscopy coupled with immunocytochemistry demonstrates that only S1 of hemp phloem fibers, both primary and secondary, is labeled by antixylan antibody [38]. The rest of the thickened cell wall does not contain epitopes for antixylan antibodies (Figure 6). This is very distinct with xylem cells, which are heavy labeled by these antibodies throughout all secondary cell wall layers [37].
Figure 5.
(A) Scheme of the sequential deposition of cell wall layers during thickening of primary and secondary phloem fibers of hemp stem. Primary and secondary fibers differ significantly in diameter as well as in the ratio of cell wall layers. The thickness of the primary cell wall and middle lamellae (ML + PCW) is comparable in primary and secondary fibers. Later deposited layer of secondary cell wall (SCW) is more developed in secondary fibers and exceeds that of primary fibers in width roughly twofold. Thеn, newly deposited layer of the tertiary (gelatinous) cell wall (Gn) is formed, which is later transformed into mature layer of tertiary (gelatinous) cell wall (G). In mature fibers, the layer Gn can be absent. (B) Cross-section of hemp secondary phloem fibers with different cell wall layers labeled with antibodies N1 raised against flax fiber-specific β-galactosidase [39]; label is absent in SCW, but present in G and especially Gn layers. (C) Primary and secondary phloem fibers on the cross-section of hemp stem stained with Calcofluor White, under UV light (lignified layers of cell walls look yellow due to lignin autofluorescence, nonlignified cell wall layers look blue); only outer layers of fiber cell wall are lignified. Bar = 1 μm (B) and 20 μm (C).
Figure 6.
Scheme illustrating the differences in distribution of polymers between cell wall layers in hemp fiber. Occurrence of cell wall epitopes for carbohydrate-binding module CBM3, antibodies LM10, LM11, RU2, N1, and JIM14, and localization of lignin deposition in various layers of phloem fiber cell wall in hemp stem.
The difference between S1 and the other layers of thickened cell wall is also obvious after labeling with antibody against fiber-specific galactosidase: S1 is not labeled, while the epitope is quite abundant in the inner two layers (Figure 5B). The antibody was raised against the enzyme isolated from flax, but it also binds cell wall in hemp phloem fibers, same as G-layers of tension wood fibers in poplar [39]. This tissue- and stage-specific β-1,4-galactosidase is necessary for maturation of cell wall structure in flax fibers, and is involved in partial trimming off the β-1,4-galactan side-chains from the backbone of rhamnogalacturonan I [40]. Similar polymer—rhamnogalacturonan I with β-1,4-galactan side-chains is also present in hemp fibers [38, 41]. Same as in flax [42, 43], fraction of this polymer is so tightly retained by cellulose that can be obtained only after complete cellulose degradation [38]. The antibody RU2 against rhamnogalacturonan I backbone [44] does not recognize any epitopes within S1 layer, but binds to the thick inner cell wall layers of hemp fibers, both primary and secondary, same as to the primary cell wall/middle lamellae region [38] (Figure 6). Cytochemical staining for pectin is also positive in the inner layers of hemp fibers [6]. Presence of acidic component indicates that inner layer of hemp fibers is similar to G-layers of tension wood and flax fibers [45, 46]. Notably, hemp fiber cell wall is not labeled by LM5 antibody specific for β-1,4-galactan [37, 38], despite the fact that the presence of corresponding polymer is biochemically proven [38, 41].
Difference between S1 and the rest of the thickened cell wall layers is further evidenced by the character of hemp fiber lignification. As usual for secondary cell wall layers, S1 gets lignified, especially at an advanced stage of hemp fiber development (Figures 5C and 6). Lignification occurs only in the outer cell wall layers of hemp fibers (middle lamellae, primary cell wall and S1): autofluorescence that is characteristic of lignin under UV light (Figure 5C), and staining for lignin by specific dyes like phloroglucinol are not observed in the thick inner layers, though they are obvious for S1 [6, 15]. The rest of cell wall remains nonlignified even in the fully formed fibers, which is especially obvious for primary fibers. Since the proportion of S1 layer in total cell wall is higher in secondary fibers than in primary ones, the degree of their lignification is higher [6, 15, 38]. This makes secondary fibers more coarse and rigid, and is often considered as the major reason for their lower quality [9].
The main component of the inner layers of cell wall is cellulose. This is evidenced by binding of CBM3—carbohydrate-binding module specific for crystalline cellulose [37] and also by high content of cellulose in hemp fibers [2, 3]. The content of hemicelluloses in hemp fibers is reduced at advanced stages of hemp fiber development [15], which may be due to the increasing proportion of cellulose-enriched inner cell wall layers. The average microfibril angle (MFA) toward the longitudinal fiber axis in hemp fibers is low; in the major cell wall layer (G/S2), it constitutes an average of 2.65° [47], meaning that the orientation of all microfibrils is close to axial. In S1, MFA is over 80° [47].
Same as in flax phloem fibers, cell wall of hemp fibers is a dynamic structure with intensive post-synthetic modifications of the deposited RG-I (Figure 5). The newly deposited portions, designated as Gn look rather loosened and contain larger amount of electron-dense material than the mature G-layer. Gn is a transient layer and is transformed into G in the course of fiber development, while the new portions of Gn are deposited (Figure 5A) [38]. The transition of Gn- into G-layer is coupled to the action of fiber-specific galactosidase [40] (Figure 5).
Together with the cell wall polymers discussed above (xylan, lignin, rhamnogalacturonan I with β-1,4-galactan side chains), immunodot analysis of isolated cell wall constituents reveals other polysaccharides, namely glucomannans, polygalacturonic acid, arabinogalactan proteins, and some xyloglucans [38]. These polymers are unevenly distributed between cell wall layers. For instance, arabinogalactan proteins are mainly detected at the inner surface of fiber cell wall, close to plasma membrane (Figure 6) [37]. The structural peculiarities of these polymers in hemp fibers, same as the possible differences in their structure in fibers of contrast quality are still to be characterized.
Absence or low content of xylan and lignin, axial orientation of cellulose microfibrils, presence of pectic components, processes of cell wall maturation with the involvement of tissue- and stage-specific galactosidase make to consider that the overall structure of hemp fibers resembles that of tension wood fibers and of flax fibers [48, 49], and that the inner layers of thickened cell wall may be viewed as tertiary cell wall. Tertiary cell wall, also named G-layer, is a fiber-specific cell wall type [19]. Its structure is based on the entrapment of RG-I by laterally interacting cellulose microfibrils. Tertiary cell walls are formed in fibers of various plant species and in many ecophysiological situations [50]. Secondary and tertiary cell walls have different mechanical properties, such as the lignified secondary cell wall provides rigidity, while tertiary cell wall adds flexibility due to the tension of cellulose microfibrils. The selection of fiber crops, like hemp, flax, ramie, has led to the extreme development of tertiary cell walls in fibers of their stems.
Summarizing, the major revealed differences between thickened cell wall of primary and secondary hemp fibers lay in the total cell wall width and in the proportion of S1 layer relative to the rest of cell wall. In secondary fibers, the cell wall width is considerably lower than in primary ones, while the proportion of S1 layer is higher. Since it is mainly S1 layer that gets lignified, secondary fibers have higher lignin content and due to that they are coarser than primary fibers. Importantly, the nanomechanical properties of tertiary cell wall as such are similar in primary and secondary fibers, as revealed by peak-force quantitative nanomechanical property mapping (PF-QNM) and micro tomography [31]. The major parameters of cell wall thickening that influence the yield and quality of hemp fibers are (1) the amount of the deposited cell wall (cell wall thickness), (2) the ratio between the cell wall and cell lumen on the fiber cross-section, (3) the proportion of S1 layer in total cell wall thickness, and (4) the set and peculiarities of structure of cell wall polymers.
Acknowledgments
We thank the colleagues from the Chuvash Research Institute of Agriculture (Tsivilsk, Russia): Dr. G.S. Stepanov, Dr. I.V. Romanova, and N.A. Trofimova—for growing the hemp plants. We also thank Dr. Paul Knox (University of Leeds) for the provided LM5 and LM11 antibodies and Dr. Fabienne Guillon (INRA, France) for the provided RU2 antibody. The research on the topic of the chapter was supported by the Russian Science Foundation, project 16-14-10256 (TG, NI, TC), and by the Russian Foundation for Basic Research, projects 15-04-02560 (PM) and 15-04-05721 (TC, MA).
\n',keywords:"plant fibers, gelatinous fibers, hemp, intrusive growth, plant cell wall, rhamnogalacturonan I",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/57169.pdf",chapterXML:"https://mts.intechopen.com/source/xml/57169.xml",downloadPdfUrl:"/chapter/pdf-download/57169",previewPdfUrl:"/chapter/pdf-preview/57169",totalDownloads:947,totalViews:485,totalCrossrefCites:2,totalDimensionsCites:2,hasAltmetrics:1,dateSubmitted:"April 26th 2017",dateReviewed:"September 13th 2017",datePrePublished:"December 20th 2017",datePublished:"May 2nd 2018",dateFinished:null,readingETA:"0",abstract:"Plant fibers in general and hemp fibers in particular have great prospects for their use in various innovative applications such as ecological, biodegradable, and renewable resources with unique properties. Such properties together with the increased strength due to high-cellulose content and specific morphological parameters are widely used to produce plant fiber–based plastic composites. The properties of plant fibers that may influence the properties of composites depend on crop processing, but the basis for them is provided during fiber development in planta. It is known that two types of bast fibers are developed in the hemp stem: primary fibers formed from procambium cells and secondary fibers that originate as a result of cambium activity. Both types of fibers may significantly vary in their yield and quality depending on the variety and growth conditions. Differences in the anatomical and morphological characteristics of the two types of hemp fibers, together with peculiarities in the composition and architecture of cell wall, influence the technical parameters of the raw material quality. Based on our study of both primary and secondary fiber development in hemp stem that was focused on the two key stages, intrusive elongation and deposition of thick cell wall layers, we suggest the set of parameters that can influence the quality of the mature fibers and trace their biological origin.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/57169",risUrl:"/chapter/ris/57169",book:{slug:"natural-and-artificial-fiber-reinforced-composites-as-renewable-sources"},signatures:"Chernova Tatyana, Mikshina Polina, Salnikov Vadim, Ageeva\nMarina, Ibragimova Nadezda, Sautkina Olga and Gorshkova\nTatyana",authors:[{id:"158372",title:"Dr.",name:"Tatyana",middleName:null,surname:"Chernova",fullName:"Tatyana Chernova",slug:"tatyana-chernova",email:"chernova.t@mail.ru",position:null,institution:null},{id:"209953",title:"Prof.",name:"Tatyana",middleName:null,surname:"Gorshkova",fullName:"Tatyana Gorshkova",slug:"tatyana-gorshkova",email:"gorshkova@kibb.knc.ru",position:null,institution:{name:"Institute of Bioorganic Chemistry",institutionURL:null,country:{name:"Russia"}}},{id:"209955",title:"Dr.",name:"Polina",middleName:null,surname:"Mikshina",fullName:"Polina Mikshina",slug:"polina-mikshina",email:"p.mikshina@gmail.com",position:null,institution:null},{id:"209956",title:"Dr.",name:"Marina",middleName:null,surname:"Ageeva",fullName:"Marina Ageeva",slug:"marina-ageeva",email:"mageeva58@mail.ru",position:null,institution:null},{id:"209957",title:"MSc.",name:"Olga",middleName:null,surname:"Sautkina",fullName:"Olga Sautkina",slug:"olga-sautkina",email:"semargla.zhar@gmail.com",position:null,institution:null}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Origin of primary and secondary fibers in the hemp stem",level:"1"},{id:"sec_3",title:"3. Intrusive elongation of primary and secondary fibers",level:"1"},{id:"sec_4",title:"4. Cell wall thickening",level:"1"},{id:"sec_5",title:"Acknowledgments",level:"1"}],chapterReferences:[{id:"B1",body:'Pickering KL, Aruan Efendy MG, Le TM. A review of recent developments in natural fibre composites and their mechanical performance. Composites Part A: Applied Science and Manufacturing. 2016;83:98-112. DOI: org/10.1016/j.compositesa.2015.08.038'},{id:"B2",body:'McDougall GJ, Morrison IM, Stewart D, Weyers JDB, Hillman JR. Plant fibres: Botany, chemistry and processing for industrial use. Journal of the Science of Food and Agriculture. 1993;62:1-20. DOI: 10.1002/jsfa.2740620102'},{id:"B3",body:'Bonatti PM, Ferrari C, Focher B, Grippo C, Torri G, Cosentino C. 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Oxford: Pergamon Press; 1990. 588 p'},{id:"B12",body:'Gorshkova TA, Sal’nikov VV, Chemikosova SB, Ageeva MV, Pavlencheva NV, Van Dam JEG. The snap point: A transition point in Linum usitatissimum bast fiber development. Industrial Crops and Products. 2003;18:213-221. DOI: 10.1016/S0926-6690(03)00043-8'},{id:"B13",body:'Sengloung T, Kaveeta L, Müssig J. Physical properties of traditional Thai hemp fiber (Cannabis sativa L.). Journal of Industrial Hemp. 2008;13(1):20-36. DOI: 10.i080/15377880801898709'},{id:"B14",body:'Chernova TE, Ageeva MV, Chemikosova SB, Gorshkova TA. The formation of primary and secondary fibers in hemp. Bulletin of the All-Russian Scientific Research Institute of Bast Crops Processing. 2005;2:6-13'},{id:"B15",body:'Crônier D, Monties B, Chabbert B. Structure and chemical composition of bast fibers isolated from developing hemp stem. Journal of Agricultural and Food Chemistry. 2005;53:8279-8289. 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Plant Physiology. 1986;82:1153. DOI: org/10.1104/pp.82.4.1153'},{id:"B21",body:'Lev-Yadun S. Intrusive growth—The plant analog of dendrite and axon growth in animals. New Phytologist. 2001;150:508-512. DOI: 10.1046/j.1469-8137.2001.00143.x'},{id:"B22",body:'Snegireva AV, Ageeva MV, Amenitskii SI, Chernova TE, Ebskamp M, Gorshkova TA. Intrusive growth of sclerenchyma fibers. Russian Journal of Plant Physiology. 2010;57:342-355. DOI: org/10.1134/S1021443710030052'},{id:"B23",body:'Garcia-Jaldon C, Dupeyre D, Vignon MR. Fibers from semi-retted hemp bundles by steam explosion treatment. Biomass and Bioenergy. 1998;14(3):251-260. DOI: 10.1016/S0961-9534(97)10039-3'},{id:"B24",body:'Sankari HS. Comparison of bast fibre yield and mechanical fibre properties of hemp (Cannabis sativa L.) cultivars. Industrial Crops and Products. 2000;11:73-84. DOI: 10.1016/S0926-6690(99)00038-2'},{id:"B25",body:'Marrot L, Lefeuvre A, Pontoire B, Bourmaud A, Baley C. Analysis of the hemp fiber mechanical properties and their scattering (Fedora 17). Industrial Crops and Products. 2013;51:317-327. DOI: org/10.1016/j.indcrop.2013.09.026'},{id:"B26",body:'Ageeva MV, Petrovská B, Kieft H, Sal’nikov VV, Snegireva AV, van Dam JEG, van Veenendaal WLH, Emons AMC, Gorshkova TA, van Lammeren AAM. Intrusive growth of flax phloem fibers is of intercalary type. Planta 2005;222(4):565-574. DOI: 10.1007/s00425-005-1536-2'},{id:"B27",body:'Akin DE. Linen most useful: perspectives on structure, chemistry, and enzymes for retting flax. ISRN Biotechnology. 2013;186534:2013. DOI: doi.org/10.5402/2013/186534'},{id:"B28",body:'Carpita N, McCann M. The cell wall. In: Buchanan BB, Wilhelm G, Jones RL, editors. Biochemistry and Molecular Biology of Plants. Rockville: American Society of Plant Physiologists; 2000. p. 52-108'},{id:"B29",body:'Gorshkova TA, Salnikov VV, Pogodina NM, Chemikosova SB, Yablokova EV, Ulanov AV, Ageeva MV, van Dam JEG, Lozovaya VV. Composition and distribution of cell wall phenolic compounds in the flax (Linum usitatissimum L.) stem tissues. Annals of Botany (Lond). 2000;85:477-486. DOI: org/10.1006/anbo.1999.1091'},{id:"B30",body:'Chemikosova SB, Pavlencheva NV, Gur’yanov OP, Gorshkova TA. The effect of soil drought on the phloem fiber development in long-fiber flax. Russian Journal of Plant Physiology. 2006;53(5):656-662. DOI: org/10.1134/S1021443706050098'},{id:"B31",body:'Bourmaud A, Malvestio J, Lenoir N, Siniscalco D, Habrant A, King A, Legland D, Baley C, Beaugrand J. Exploring the mechanical performance and in-planta architecture of secondary hemp fibres. Industrial Crops and Products. 2017;108:1-5. DOI: org/10.1016/j.indcrop.2017.06.010'},{id:"B32",body:'Snegireva AV, Ageeva MV, Vorob’ev VN, Anisimov AV, Gorshkova TA. Plant fiber intrusive growth characterized by NMR method. Russian Journal of Plant Physiology. 2006;53:163-168. DOI: org/10.1134/S1021443706020038'},{id:"B33",body:'Behr M, Legay S, Žižková E, Motyka V, Dobrev PI, Hausman J-F, Lutts S, Guerriero G. Studying secondary growth and bast fiber development: the hemp hypocotyl peeks behind the wall. Frontiers in Plant Science. 2016;7:1733. DOI: 10.3389/fpls.2016.01733'},{id:"B34",body:'Guerriero G, Behr M, Legay S, Mangeot-Peter L, Zorzan S, Ghoniem M, Hausman J-F. Transcriptomic profiling of hemp bast fibres at different developmental stages. Scientific Reports. 2017;7:4961. DOI: 10.1038/s41598-017-05200-8'},{id:"B35",body:'Mokshina N, Gorshkov O, Ibragimova N, Chernova T, Gorshkova T. Cellulosic fibres of flax recruit both primary and secondary cell wall cellulose synthases during deposition of thick tertiary cell walls and in the course of graviresponse. Functional Plant Biology. 2017;44(8):820-831. DOI: org/10.1071/FP17105'},{id:"B36",body:'McCartney L, Marcus SE, Knox JP. Monoclonal antibodies to plant cell wall xylans and arabinoxylans. The Journal of Histochemistry and Cytochemistry. 2005;53:543-546. DOI: 10.1369/jhc.4B6578.2005'},{id:"B37",body:'Blake AW, Marcus SE, Copeland JE, Blackburn RS, Knox JP. In situ analysis of cell wall polymers associated with phloem fibre cells in stems of hemp, Cannabis sativa L. Planta. 2008;228:1-13. DOI: org/10.1007/s00425-008-0713-5'},{id:"B38",body:'Gorshkova TA, Gurjanov OP, Mikshina PV, Ibragimova NN, Mokshina NE, Salnikov VV, Ageeva MV, Amenitskii SI, Chernova TE, Chemikosova SB. Specific type of secondary cell wall formed by plant fibers. Russian Journal of Plant Physiology. 2010;57(3):328-341. DOI: org/10.1134/S1021443710030040'},{id:"B39",body:'Mokshina NE, Ibragimova NN, Salnikov VV, Amenitskii SI, Gorshkova TA. Galactosidase of plant fibers with gelatinous cell wall: identification and localization. Russian Journal of Plant Physiology. 2012;59(2):246-254. DOI: 10.1134/S1021443712020082'},{id:"B40",body:'Roach MJ, Mokshina NY, Badhan A, Snegireva AV, Hobson N, Deyholos MK, Gorshkova TA. Development of cellulosic secondary walls in flax fibers requires beta-galactosidase. Plant Physiology. 2011;156(3):1351-1363. DOI: 10.1104/pp.111.172676'},{id:"B41",body:'Vignon MR, Garcia-Jaldon. Structural features of the pectic polysaccharides isolated from retted hemp bast fibres. Carbohydrate Research. 1996;296:249-260. DOI: 10.1016/S0008-6215(96)00226-1'},{id:"B42",body:'Gurjanov OP, Ibragimova NN, Gnezdilov OI, Gorshkova TA. Polysaccharides, tightly bound to cellulose in the cell wall of flax bast fibre: Isolation and identification. Carbohydrate Research. 2008;72:719-729. DOI: 10.1016/j.carbpol.2007.10.017'},{id:"B43",body:'Mikshina PV, Gurjanov OP, Mukhitova FK, Petrova AA, Shashkov AS, Gorshkova TA. Structural details of pectic galactan from the secondary cell walls of flax (Linum usitatissimum L.) phloem fibres. Carbohydrate Polymers. 2012;87:853-861. DOI: 10.1016/j.carbpol.2011.08.068'},{id:"B44",body:'Ralet M-C, Tranquet O, Poulain D, Moise A, Guillon F. Monoclonal antibodies to rhamnogalacturonan I backbone. Planta. 2010;231:1373-1383. DOI: 10.1007/s00425-010-1116-y'},{id:"B45",body:'Bowling AJ, Vaughn KC. Immunocytochemical characterization of tension wood: Gelatinous fibers contain more than just cellulose. American Journal of Botany. 2008;95:655-663. DOI: 10.3732/ajb.2007368'},{id:"B46",body:'Gorshkova TA, Gurjanov OP, Mikshina PV, Ibragimova NN, Mokshina NE, Salnikov VV, Ageeva MV, Amenitskii SI, Chernova TE, Chemikosova SB. Specific type of secondary cell wall formed by plant fibers. Russian Journal of Plant Physiology. 2010;57:328-341. DOI: 10.1134/S1021443710030040'},{id:"B47",body:'Dai D, Fan M. Characteristic and performance of elementary hemp fibre. Materials Sciences and Applications. 2010;1:336-342. DOI: 10.4236/msa.2010.16049'},{id:"B48",body:'Mellerowicz EJ, Gorshkova TA. Tensional stress generation in gelatinous fibres: A review and possible mechanism based on cell-wall structure and composition. Journal of Experimental Botany. 2012;63(2):551-565. DOI: 10.1093/jxb/err339'},{id:"B49",body:'Gorshkova T, Mokshina N, Chernova T, Ibragimova N, Salnikov V, Mikshina P, Tryfona T, Banasiak A, Immerzeel P, Dupree P, Mellerowicz EJ. Aspen tension wood fibers contain β-(1-4)-galactans and acidic arabinogalactans retained by cellulose microfibrils in gelatinous walls. Plant Physiology. 2015;169(3):2048-2063. DOI: 10.1104/pp.15.00690'},{id:"B50",body:'Mikshina P, Chernova T, Chemikosova S, Ibragimova N, Mokshina N, Gorshkova T. Cellulosic Fibers: role of matrix polysaccharides in structure and function. In: van de Ven T, Godbout L, editors. Cellulose – Fundamental Aspects. Rijeka: InTech; 2013. p. 91-112. DOI: 10.5772/51941'}],footnotes:[],contributors:[{corresp:null,contributorFullName:"Chernova Tatyana",address:null,affiliation:'
Interdisciplinary Center for Analytical Microscopy, Kazan Federal University, Kazan, Russia
Interdisciplinary Center for Analytical Microscopy, Kazan Federal University, Kazan, Russia
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\n
1. Introduction
\n
Mosquitoes are small, midge-like flies that constitute the family Culicidae. Females of most species are ectoparasites feeding on vertebrates’ blood through piercing the hosts’ skin to suck the blood. To-date, approximately 3500 species of the Culicidae have been described. The family Culicidae is a large and abundant group which occurs throughout temperate and tropical regions of the world and well beyond the Arctic Circle [1]. There are two subfamilies of Culicidae, that is, the Anophelinae (3 genera) and the Culicinae (110 genera). The subfamily Culicidae, Aedes is the largest tribe of mosquitoes with 1256 species classified into 10 genera: Aedes sensu (931), Armigeres (58), Eretmapodites (48), Haemagogus (28), Heizmannia (38), Opifex (2), Psorophora (49), Udaya (3), Verrallina (95), and Zeugnomyia (4) [2].
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The public health concern of Aedes mosquitoes particularly Ae. aegypti and Ae. albopictus in the transmission of arboviruses such as dengue virus, chikungunya virus, ZIKV virus, and yellow fever virus is kept on increasing globally. Over half of the world’s population is at risk of dengue and chikungunya infections [3]. The Caribbean, South America, and Europe are no longer spared from chikungunya infection, a disease which was previously limited to Africa and Asia [3]. According to the World Health Organization, about 2.5 billion people globally live in dengue endemic regions [4]. Dengue is the most worldwide important mosquito-transmitted viral infection [4]. Over 100 countries in Africa, North and South America, Southeast Asia, Europe, and the Pacific are reported to have had severe dengue outbreaks [5]. The annual occurrence of dengue fever infections ranges from 50 to 100 million with which around 500,000 facing severe morbidity causing to over 20,000 mortalities, pediatrics beings the most cases [5]. The chikungunya virus infections (CHIKV) have been documented in over 60 countries in Asia, Africa, Europe, and the Americas [6]. The estimated number of chikungunya cases in Americas in 2016 was 693,000, and Zika virus (ZIKV) disease was 500,000 [6, 7]. Yellow fever cases in Africa were 130,000 with an estimated 31,000 annual disability adjusted life years and 500 deaths [8, 9].
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About 80% of the world’s population is at risk for at least of exposure to one vector-borne disease; these diseases account for about 17% of the estimated global burden of communicable diseases and cause over 700,000 deaths annually, affecting disproportionately poorer populations [6, 9]. They hamper economic development through direct medical costs and indirect costs such as the loss of productivity and tourism. The social, demographic, and environmental factors strongly influence transmission patterns of vector-borne pathogens. Vector control is an important component for decision science in the prevention and control of vector-borne disease approaches. Consequently, the global distribution and ecology of these vectors and the geographical determinants of their ranges are essential in order to be effective. Therefore, it is important to work out where these mosquito species are found around the globe to identify the areas at risk. It is also important to predict where these species could become established if they were introduced, in order to identify areas that could become at risk in the future.
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1.1. Aedes distribution
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Ae. aegypti and Ae. albopictus are worldwide distributed between 35° N and 35° S, latitudes that roughly correspond to a 10°C winter isotherm which appears to be the limiting temperatures that the species can tolerate while overwintering [5]. The species are highly adapted to urban environments, breeding in stagnant water found in manufactured containers, garbage heaps, and tyres. However, the distribution of Ae. albopictus has been highly biased to temperate climates [10] though the vector is now widely distributed throughout the Americas (excluding Canada), Europe, Asia, Africa, Australia, and the Pacific [11].
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The geographic distribution of Ae. aegypti based on the order of higher levels of occurrence for each continent reveals that in the Americas, the Brazil ranks the highest (Table 1). In Africa, occurrence of the vectors have been recorded in Senegal, Cameroon, Kenya, Tanzania, Ivory Coast, Nigeria, Madagascar, Gabon, and Sierra Leone (Table 1). In Asia/Oceania, occurrence of Ae. aegypti has been reported and documented (Table 1) [3, 12].
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Country
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Occurrences
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Country
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Occurrences
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Country
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Occurrences
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\n\n\n
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Ae. aegypti
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Americas
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Brazil
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5044
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Europe/Africa
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Senegal
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112
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Asia/Oceania
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Taiwan
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9490
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USA
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436
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Cameroon
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55
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Indonesia
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603
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Mexico
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411
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Kenya
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52
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Thailand
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495
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Cuba
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177
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United Republic of Tanzania
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44
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India
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423
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Argentina
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170
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Côte d’Ivoire
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40
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Australia
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282
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Trinidad and Tobago
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152
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Nigeria
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35
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Viet Nam
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223
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Venezuela
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130
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Madagascar
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28
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Malaysia
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112
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Colombia
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128
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Gabon
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27
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Singapore
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44
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Puerto Rico
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120
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Mayotte
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20
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Philippines
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36
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Peru
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89
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Sierra Leone
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20
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Cambodia
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29
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Ae. albopictus
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Americas
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Brazil
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3441
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Europe/Africa
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Italy
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203
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Asia/Oceania
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Taiwan
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15,339
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USA
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1594
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Madagascar
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58
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Malaysia
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186
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Mexico
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50
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Cameroon
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42
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Indonesia
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161
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Cayman Islands
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15
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France
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37
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India
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150
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Haiti
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13
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Gabon
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27
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Japan
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97
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Guatemala
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12
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Albania
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22
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Thailand
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82
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Venezuela
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7
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Mayotte
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21
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Singapore
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44
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Colombia
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3
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Greece
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18
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Lao People’s Democratic Republic
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26
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Cuba
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3
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Israel
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17
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Philippines
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22
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Puerto Rico
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3
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Lebanon
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15
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Viet Nam
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18
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Table 1.
The Aedes aegypti and Ae. albopictus distribution globally.
Note: This table was contributed by Kramer a leading author of the paper published in E-life journal (https://doi.org/10.7554/eLife.08347.003).
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1.2. Ecology of Aedes mosquitoes
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1.2.1. Aedes aegypti
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Ae. aegypti is an arthropod closely associated with humans and their habitats. They are mostly anthropophilic [13] with high preference to the urban environment [14]. They get blood meals from human, and human creates conducive environment for their population growth through up haphazardly disposal of water-holding containers/obsoletes around our homes. The mosquito lays her eggs on the sides of containers with water, and eggs hatch into larvae after a rain or flooding. A larva changes into a pupa in about a week and into a mosquito in 2 days. The Aedes main habitat is aquatic, and they can thrive better from tree cavities to toilets. People also furnish shelter as Ae. aegypti preferentially rests in darker cool areas, such as closets leading to their ability to bite indoors.
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Ae. aegypti has adaptations to the environment that makes them highly resilient, or with the ability to rapidly bounce back to initial numbers after disturbances resulting from natural phenomena (e.g., droughts) or human interventions (e.g., control measures). One such adaptation is the ability of the eggs to withstand desiccation (drying) and to survive without water for several months on the inner walls of containers. For example, if we were to eliminate all larvae, pupae, and adult Ae. aegypti at once from a site, its population could recover 2 weeks later as a result of egg hatching following rainfall or the addition of water to containers harboring eggs.
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It is likely that Ae. aegypti is continually responding or adapting to environmental change. For example, it was recently found that Ae. aegypti is able to undergo immature development in broken or open septic tanks resulting in the production of hundreds or thousands of Ae. aegypti adults per day. In general, it is expected that control interventions will change the spatial and temporal dispersal of Ae. aegypti and perhaps the pattern of habitat utilization.
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1.2.2. Aedes albopictus
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Aedes albopictus (Stegomyia albopicta), from the mosquito (Culicidae) family, also known as (Asian) tiger mosquito or forest mosquito, is a mosquito native to the tropical and subtropical areas of Southeast Asia; however, in the past few decades, this species has spread to many countries through the transport of goods and international travel [15]. The eggs of Ae. albopictus are desiccation resistant, which enhance survival in inhospitable environments [16]. Ae. albopictus is among the aggressive outdoor species of mosquito, and they are day biter that has a very broad host range and attacks humans, livestock, amphibians, reptiles, and birds [17]. Their biting rate level can be as high as 30 to 48 bites per hour [18]. Ae. albopictus survives at a large range of temperatures [19].
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Ae. albopictus is a treehole mosquito, and so its breeding places in nature are small, restricted, shaded bodies of water surrounded by vegetation. It inhabits densely vegetated rural areas. However, its ecological flexibility allows it to colonize many types of man-made sites and urban regions. It may reproduce in cemetery flowerpots, birdbaths, soda cans and abandoned containers, and water recipients. Tyres are particularly useful for mosquito reproduction as they are often stored outdoors and effectively collect and retain rainwater for a long time. The addition of decaying leaves from the neighboring trees produces chemical conditions similar to tree holes, which provides an excellent substrate for breeding. Ae. albopictus can also establish and survive throughout nonurbanized areas lacking any artificial containers, raising additional public health concerns for rural areas [17].
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1.2.3. Aedes mosquitoes life cycle
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Aedes mosquito species, Ae. aegypti, and Ae. albopictus are major public health concern due to their role in transmission of diseases [3]. Ae. aegypti mosquito is widespread in (sub-)tropical regions and is largely responsible for vector-borne arboviral infections, yellow fever virus (YFV), ZIKV, dengue virus (DENV), West Nile virus (WNV), CHIKV and transmission, and outbreaks in various regions [3, 7]. The Ae. aegypti is known to have high vectorial capacity due to its anthropophilic behavior, well domesticated, and adapted to survive in different geographical regions including Africa, Americas, Asia, and Europe [1, 3].
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The Aedes spp. mosquitoes are known to have a complex life cycle involving aquatic and terrestrial life [2]. Mosquitoes acquire the infection after a blood-meal form the host in order for the eggs to develop. The vector needs water to lay their eggs in the preferred breeding container, including tyres, water storage containers, disposed tyres, coconut shells, and flowerpots [20]. Aedes spp. prefers to lay their eggs on the inner wet walls of containers with water, hence the name “container breeder”. The development of the eggs occurs between 2 and 7 days in the aquatic phase (Figure 1) where the larvae hatch from the eggs. The larva survival depends on the microorganisms found in the aquatic environment. Larvae go through developmental stages (stage 1–4) in which they molt or shed their skin; these larval stages are called the first to fourth instars [20]. When a larva is a fully grown fourth instar, it undergoes metamorphosis into a new form called a pupa in approximately 4 days, the “cocoon” stage for the mosquito. This developmental stage of the mosquitoes also occurs in the aquatic environment. After 1–2 days, the fully developed adult mosquito forms and breaks through the skin of the pupa and a fully grown adult emerges. The adult mosquito is able to fly and has a terrestrial habitat inhabiting inside and outside households [20].
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Figure 1.
Aedes mosquito life cycle in aquatic and terrestrial phases.
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Interestingly, Aedes has developed a survival mechanism during the dry seasons; the eggs can enter a dormancy (quiescence) for up to 8 months at the end of embryogenesis [21]. If the habitat is dry, the eggs remain dormant but after rainfall, the eggs hatch and development continues [20]. In addition to being desiccation resistant, Aedes spp. is well adapted to produce, eggs can withstand months of dormancy, so-called “extended quiescence” in the unfavorable abiotic environment [21]. The male Aedes spp. mosquitoes feed on flowers’ nectar or plant juices, unlike the female that needs a blood meal [22]. The vector becomes infected when they feed on infected humans, and transmission may occur when the vector bites the host, which is believed to be promoted by mosquito salivary protein.
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Historically, Ae. aegypti is believed to have originated from zoophilic subspecies Ae. Aegypti formosus inhabiting forests in sub-Saharan Africa [12]. This subspecies is found in the forests, breed in the tree holes and feeding on other mammals. The evolution of the ancestral Ae. aegypti resulted in the domesticated Ae. aegypti subspecies with a strong preference for biting humans and breed in man-made containers [20]. This evolved as the dominant vector of several diseases including yellow fever and DENV, ZIKV infections worldwide. The domestication of the vector was associated with the human migrations, trade, transportation, and urbanization [20, 23]. The domestic Ae. aegypti thrive in (sub-)tropical and temperate regions and can inhabit either terrestrial or aquatic depending on the stages of the growth. Ae. aegypti is primarily a container breeding vector and is known to predominate in urban areas where there is the vast composition of favorable man-made breeding container environment [20]. The breeding sites range from natural to artificial including vegetation, discarded tyres, discarded containers, bottle tops, water storage containers (especially in places with erratic tap water supply), flowerpots and vases, metal drums, and coconut shells [9, 24]. Other breeding sites include the open or unsealed septic tanks, water wells, and water meters. The ecological factors determine the crucial characteristics of different stages of the life and eventually its success. The Ae. aegypti larvae feed on nutritious materials available in the aqueous phase in the breeding containers including the plant particles, animal debris, and phytoplankton such as microalgae found in the water-filled containers [25]. The ecological characteristics are important in the life cycle of the adult vector including the longevity, fecundity body mass, and vectorial competence [26]. For instance, some algae species, Cladophora sp., Chlorella ellipsoidea, and Rhizoclonium hieroglyphicum, were shown to exhibit larvicidal properties that affect the development of the immature stages [25]. Evidence suggests that the developmental stage from first instar larval stage to adult mosquito is faster when the organic matters are abundant in the breeding container, in addition, the survival rate of the immature stage is enhanced [27]. In contrast, low concentration or exhaustion of the nutrients is required to trigger pupation presumably in response to the increasing level of ecdysteroid hormone [3, 28, 29]. Temperature is important for the survival larva density and competence of the Ae. aegypti. In areas where the temperature is warmer, the development of the aquatic stage temperature was associated with shorter development time from hatch to the emergence of the adult mosquito [4]. Similarly, longer light exposure was also shown to shorten the development time [30]. The evidence explains the widespread distribution and pattern of Ae. aegypti in (sub-)tropical regions. Furthermore, evidence suggests increasing Ae. aegypti abundance in urban areas leading to outbreaks [31]. It is evident that developing countries are becoming more urbanized; however, poor city planning and sanitation have increased mosquito breeding sites [7]. The “ecological plasticity” exhibited by the vector is arguably among the reasons for reason that explain it its worldwide widespread and success as a human vector.
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1.3. Insecticide resistance in Aedes spp.
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The emergence of insecticide resistance to multiple classes of insecticides has been widely reported in Ae. aegypti in different regions [24, 32, 33, 34]. WHO defines resistance as the ability of mosquitoes to survive exposure to a standard dose of insecticide; this ability may be the result of physiological or behavioral adaptation [35]. The emergence and spread of resistance to the main insecticides could compromise the effectiveness of the preventive measures, operational implementation of control programs, and outbreak management.
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1.4. Mechanisms of insecticide resistance
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There are three major categories of insecticide resistance that have been described, namely, physiological resistance (target-site resistance and metabolic resistance) and behavioral avoidance. First, physiological resistance may develop due to the target-site resistance. Target site mutations are known to cause amino acid substitutions, which could affect the influx of insecticides into the target site. This may compromise the action of the insecticide rendering the vector tolerant or fully resistant to the insecticide. Another form of physiological resistance is due to metabolic resistance due to detoxification of insecticides by cytochrome P450 monooxygenases which allow the resistant vector to metabolize insecticides [36]. Glutathione S-transferases (GSTs) and carboxylesterases (ESTs) are also described in this process. Over expression of P450s was associated with insecticide resistance in diverse vector species including Ae. aegypti [37]. The resistant vectors accumulate high levels of efficient enzymes that detoxify the toxins. The second mechanism of resistance is known as behavioral adaptation or avoidance of the vector, this is well characterized in Anopheles mosquitoes. Therefore, monitoring insecticide resistance is crucial in the implementation of vector control strategies.
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1.4.1. Physiological resistance in Aedes spp.
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In Tanzania, like many other African settings, there is limited information on the Ae. aegypti resistance, most of the resistance data were collected mainly in the Americas and Asia. Our recent study in Dar es Salaam [24] demonstrated that the majority of Ae. aegypti strains were resistant to pyrethroid class of insecticide; mortality ranging from 83 to 92% in Dar es Salaam City. Data on molecular markers of resistance are scarce; however, studies elsewhere have correlated the occurrence of the knockdown resistance (kdr) mutations and resistance to pyrethroid and DDT [29, 34, 37, 38].
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The mechanism of action of the pyrethroid compounds is through their toxic effect and subsequent disruption of the VGS channels in the insect nervous system [32]. The evidence suggests that Ae. aegypti resistance to pyrethroids is conferred by the kdr mutations in the VGS channel [29, 39]. Nonsynonymous mutations in kdr gene are associated with insecticide resistance to DDT and pyrethroids on codon V1016I and F1534C in domains II and III of the VSG channel in Aedes spp. [40]. Other studies demonstrated the role of kdr gene mutation I1011M/V and F1269C in association with Ae. aegypti resistance [33, 34, 41]. In African settings, the occurrence of F1534C in concurrence with the V1016I mutation was also observed in Ghanaian Ae. aegypti population [42]. The more recent study demonstrated the significant role of kdr mutation V410L alone or in combination with the F1534C in reducing the sensitivity of Ae. aegypti to both type I (e.g., permethrin) and type II (e.g., deltamethrin) pyrethroids [32].
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In addition to the kdr mutations, metabolic resistance is also know to lead to a physiological resistance due to the increase in the synthesis of detoxifying enzymes or in their specificity to metabolize the insecticide, both resulting in an enhancement of the insect detoxifying capacity of the vector [43, 44]. The P450 monooxygenases were shown to play a significant role in modulating resistance as revealed by high-throughput assays, by comparing the overall profile at genomic and transcriptome levels between resistant and susceptible populations [37]. A study that characterized several P450s, four CYP’s, 9 J32, 9 J24, 9 J26, and 9 J28, conferring insecticide resistance in Ae. aegypti [37]. The CYPs were shown to be capable of metabolizing deltamethrin and permethrin; two common pyrethroid-based insecticides are widely used in vector interventions. Furthermore, there is evidence on the role of glutathione transferase (GST) enzymes in conferring resistance to several classes of insecticides [45]. In Ae. aegypti, the GST occurs as a cluster of genes in chromosome 2 and is shown to play a significant role in the metabolism of DDT [46]. Over expression of the GST enzyme is associated with DDT and pyrethroid resistant in Ae. aegypti populations. We, therefore, characterized additional members of this class in Ae. aegypti and provide evidence for a role of two additional GSTs in conferring resistance to insecticides.
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1.4.2. Behavioral resistance in Aedes spp.
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Thus is defined as the ability of a vector to detect and escape from an insecticide-treated area and avoid the toxin. This type of resistance has been shown in different classes of insecticides, including organochlorines, organophosphates, carbamates, and pyrethroids [47]. It has been shown that vectors are capable of avoiding feeding if they come across certain insecticides or escape the area sprayed with the insecticides. There are currently limited studies exploring this mechanism of resistance in Ae. aegypti. This paucity of information could hamper control programs since insecticide resistance could spread and render the insecticides ineffective. Therefore, more studies to assess the current susceptibility status of insecticides used for vector control are needed to describe the status to support control strategies.
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2. Disease transmission by Aedes aegypti
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Ae. aegypti mosquito is a major vector of dengue virus represented by four closely related serotypes called dengue 1, 2, 3, and 4 cause different illness including dengue fever, dengue shock syndrome, and dengue haemorrhagic fever. Dengue virus (DENV) belonging to the family Flaviviridae and genus Flavivirus [48].
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Transmission of dengue fever (DF) occurs when a female Aedes spp. mosquito obtains its blood meal from an infected person during the period of viraemia. Mosquito-borne viruses multiply in both invertebrate and vertebrate cells where they cause cytopathic effects and cell destruction. Vector mosquitoes become infected when they feed on blood of a viremic vertebrate host in which there are sufficient circulating viral particles to provide an infectious dose to the mosquito.
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A mosquito with salivary gland infection may transmit infectious virions during salivation as it probes the tissues of another vertebrate host. Transovarial transmission of virions occurs from the female mosquito to her progeny, and females of the next generation can transmit the virus orally without having been infected through blood feeding. There is also a venereal transmission of some arboviruses from male to female mosquito as observed and reported by Amarasinghe and others [49] (Figure 2).
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Figure 2.
Arboviral transmission cycle vectored by Aedes mosquitoes.
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Transmission of dengue virus occurs in 3 cycle, namely, enzootic cycle, epizootic cycle, and epidemic cycle. The enzootic cycle involves monkey-Aedes-monkey cycle, and this cycle is primitive and has been reported in South Asia and Africa [50]. The second is epizootic cycle, which involves the transmission of dengue virus from nonhuman primates to the next human in epidemic cycles by Aedes mosquito. Lastly, the epidemic cycle where the transmission cycle is through human Ae. aegypti contact, human cycle with periodic, or cyclical epidemic (Figure 2).
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In this life cycle (human-to-Ae. aegypti mosquito-to-human cycle), the main dengue virus transmission is through mosquito that usually acquires the virus after feeding on the blood of an infected person. Replication of the virus occurs in the epithelial lining of the mosquito’s midgut and then the virus move to haemocoele to infect the salivary glands. The virus can be transmitted though saliva during probing or blood feeding. The extrinsic incubation period may take 8–12 days, and this mosquito remains infected in all her life [50].
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Infected humans are the major carriers of the virus where mosquito can acquire the virus through biting. The incubation time varies from virus to virus, but generally, arboviruses exhibit between 2–15 days from inoculation to development of clinical symptoms. During this period, Aedes mosquito can acquire the virus after feeding this person.
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The reemergence of dengue disease in other places may be associated with the transovarial (via the eggs) transmission of dengue virus by Ae. aegypti. Dengue fever cannot spread directly from one person to another. Usually, Ae. aegypti prefers to feed mammalian hosts and will like to feed on humans, and even in the presence of other hosts (anthropophilic behavior), this behavior together with multiple feeding habit and highly domesticated behavior can make it an efficient vector.
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3. Seasonality and intensity of transmission
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Usually, dengue transmission occurs in rainy seasons with appropriate temperature and humidity for surviving of adult and larva mosquito. On the other hand, in arid areas, the rainfall is scant, and therefore, during the dry season, the man-made containers become the main breeding sites for the Aedes mosquito. Therefore, this can increase disease transmission.
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In the ambient temperature, the life cycle of Aedes is shortening; also, there is production of small size mosquitoes, which may lead to the reduction of extrinsic incubation period. This small size mosquito may take more blood meal for egg production, which may lead to the increase in the number of infected mosquito and speedup the disease epidemic in the next dry season [50, 51, 52].
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Several entomological factors have been associated with the initiation and maintenance of the epidemic including behavior, density, and vectorial capacity of mosquito vector population and introduction of the virus into a community.
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4. Control and surveillance
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4.1. Community education
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This can be done by professionals by giving the public awareness, which can help to empower people to take control of mosquito breedings around their surroundings and adult control. The public can be provided with the tools needed to reduce mosquito annoyance. This is when the community, families, and individuals involved in planning and implementation of local vector control activities in order to ensure that the program meets priorities and the needs of the people in the community.
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4.2. Larval mosquito control
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Frequent larval breeding sites should be searched and treated as frequent as possible by trained field technicians and trained community members. Mosquito elimination in larval stages before emerging to adults will reduce the adult mosquito population. Reduction of mosquito breeding sites such as jars, barrels, pots, vases, bottles, tins, water coolers, and tyres can be done by environmental management, removing of solid waste and managing artificial man-made habitats. All domestic water storage containers should be cleaned and covered daily.
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4.3. Adult control of Aedes aegypti
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This should aim to control Ae. aegypti population. The use of insecticides such as lambda cyhalothrin- or deltamethrin-treated material by hanging them on windows and used as water jar covers may reduce Ae. aegypti population [53]. The use of insecticide space spraying, coils, and vaporizers in the community may reduce the mosquito population.
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4.4. Use of repellents
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Application of repellents such as DEET, DIMP, and of like is of paramount importance in reducing or controlling human to vector contact. The application should be done during active hours of the day.
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4.5. Surveillance
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Surveillance is important detect mosquito species in a certain area and changes in populations. By having valuable data, we are capable of more successfully time larvicide applications and more correctly target the adulticide activities. The WHO recommends of regular household surveys of Aedes spp. collecting evidence on the ecological and epidemiological indices to guide prevention and control strategies. This involves determining the habitat productivity, preference of the breeding sites, containers for the presence of egg, larvae and pupae as well as the collection of adult mosquitoes for further identification. Larval surveys involve identifying the presence of immature mosquitoes in breeding sites such as discarded tyres, containers, and water storage vases in the defined targeted area. Through this assessment, it is possible to identify most containers that are positive for Aedes spp. and parameters such Container Index (CI) and Breteau Index (BI) [6]. On the other hand, pupal survey is performed in houses and other breeding sites to identify the productivity in the breeding habitat [7]. In addition, surveys to determine the prevalence and circulating serotypes of DENV, ZIKV, CHIKV, and YFV as a part of regular surveillance are required to inform strategies to prevent transmission and provide early warning signal of outbreaks of clinical infections.
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5. Discussion
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Ae. aegypti remains a serious public health threat due to its importance in arboviral transmission, DENV, CHIKV, and ZIKV transmission. Globally, the incidence of DENV infections is on the rise, and recently, reemergence of CHIKV and ZIKV has been observed. Vaccine, prophylaxis, and therapeutics for most arboviral infections are still in development pipeline; hence, integrated vector management remains the cornerstone to stop outbreak transmission and sustainable control. Therefore, understanding of the ecology is important for outbreak prediction and effective planning of strategies to control transmission of arboviral infections.
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Studies on the ecology of Ae. aegypti are important to better understand the preference of the vector in terms of the oviposition and colonization of mosquitoes [8]. The ecological factors play a role on influencing the population dynamics of larvae and pupae. The evidence is clear that both abiotic and biotic factors are important determinants of adulthood characteristics of life cycle such as longevity, fecundity, and body size [8, 9]. The factors are important to explain the vectorial capacity of Ae. aegypti on disease transmission. Furthermore, there is compelling evidence that Ae. aegypti is most productive in containers, which varies among regions, geographical settings, and seasonality [10]. In addition, it undoubtedly clears that water storage containers and discarded containers influence the vector density and risk of arboviral transmission particularly in poorly planned cities in (sub-)tropical regions [11]. Unless appropriate actions are taken, increasing urbanization, poor environmental management will continue to influence the stability of the Ae. aegypti populations. In addition, the vector density is influenced significantly by environment factors and urbanization [9, 11, 13]. Ae. aegypti feed exclusively on human and is increasingly a threat particularly in unplanned (peri-)urban areas. The recent data highlight the increasing Aedes spp. abundance and urbanization that could potentially escalate the risk of arboviral outbreaks [31]. Furthermore, the environment contributes to the breeding and ecological colonization of the vector. The presence of organic nutrients and microorganisms such as cyanobacteria seems to have influence on the productivity and development of Ae. aegypti [11, 15]. The presence of microalgae in the larval habitats, therefore, represents high adequacy of nutrients for immature stages of Ae. aegypti [11, 15]. Microalgae are associated with the presence and abundance of the vectors being the source of food for the larvae in breeding habitats. The evidence suggests that better understanding of these factors may be a useful indicator for mosquito population control. Measures to control microalgae to deprive nutrients to the vector could be explored for additional measures of the vector control. Importantly, approaches targeting immature stages of Ae. aegypti are highly recommended for effective and sustainable vector control. Ae. aegypti vector lays eggs in containers, buckets, care tyres and water storage vases; thus, the appropriate intervention such as proper disposal and management of containers and discarded tyres for source reduction could prove effective in reducing the vector population and mitigate the risk of arboviral transmission. Therefore, implementation of strategies to address the challenge of reemergence and expansion of arboviral infections will require a strong multisector commitment and integration for effective surveillance and control at regional, national, and program levels.
\n
There is widespread resistance to the commonly widely used insecticide, pyrethroids and organophosphates in Aedes spp. control. Insecticide resistance is likely to impact disease outbreak and transmission measures and cost of the interventions [16, 17]. This is currently a major concern in South America where organophosphates, pyrethroids, and DDT have been widely used in vector control. However, there is also evidence on decreased susceptibility to pyrethroids in Sub-Saharan Africa and Asia [10, 13]. The origin and evolution of Aedes spp. resistance to insecticides remain unclear; however, it is assumed that the use of the insecticide in other vector interventions such as malaria control and agricultural may have exerted selective pressure on the Aedes spp. The mechanism of Ae. aegypti resistance to insecticides seems to be mediated by the nonsynonymous mutations kdr gene [18, 19]. Other studies suggest the role of enhanced enzymatic biodegradation or sequestration [20, 21]. Studies suggest the potential role of the metabolic enzymes including cytochrome P450s and GSTs in conferring resistance to pyrethroids and organophosphates in Aedes spp. Evidence on the possible behavioral resistance or avoidance is patchy, and more investigation is needed to understand how it may affect the current interventions. Despite the worsening Ae. aegypti resistance to pyrethroids and organophosphates, studies on susceptibility profile of Bti are reassuring that Aedes spp. retains considerable susceptibility to the biolarvicides [17]. Therefore, Bti remains a suitable alternative for prevention and control tool in regions where resistance to pyrethroids is widespread. To mitigate the risk of resistance and its public health consequences, it is crucial to strengthen monitoring and surveillance at all levels. Susceptibility testing of the commonly used insecticides and biolarvicides using the standardized WHO bioassay protocol should be integrated as part of the surveillance program and profiling of molecular markers of resistance may be considered as appropriate.
\n
\n
\n
6. Conclusion
\n
Aedes aegypti is the most important vector in outbreak and transmission of arboviral infections. The environmental factors favor mosquitoes and risk of disease transmission. The diseases are expanding particularly in (sub-)urban settings with frequent water shortage, high human population, poor planning, and poor waste disposal systems. Primary prevention and control measures are to reduce the vector exposure, but current vector control tools are unsustainable and there is increasing threat due to insecticide resistance. Integration of Aedes spp. vector control with other ongoing program and coordination of insecticide resistance monitoring and management is crucial to increase the impact of interventions. Future interventions will require deployment of effective vaccines against arboviral infections combined with integrated vector management.
\n
\n\n',keywords:"Aedes aegypti, ecology, insecticide resistance, control, arboviruses",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/64007.pdf",chapterXML:"https://mts.intechopen.com/source/xml/64007.xml",downloadPdfUrl:"/chapter/pdf-download/64007",previewPdfUrl:"/chapter/pdf-preview/64007",totalDownloads:1287,totalViews:741,totalCrossrefCites:1,dateSubmitted:"March 20th 2018",dateReviewed:"September 10th 2018",datePrePublished:"November 5th 2018",datePublished:"January 30th 2019",dateFinished:null,readingETA:"0",abstract:"Aedes aegypti (Stegomyia) has been human vectors for many human diseases globally. In recent years, dengue virus has been diagnosed in different regions such as Asia and Latin America vectored by Aedes spp. mosquitoes. Dengue cases have been reported again in the several parts of African and other continental hospital. The different types of breeding sites have been found to be abundant in both urban and rural areas. The abundance of adult Ae. aegypti and habitat productivity in different settings escalates the risk of dengue transmission if viruses are found in asymptomatic population. The insecticide resistance has been found to occur in the wild population of Aedes aegypti to insecticides commonly used for indoor residual spray and long-lasting insecticidal net treatments. The control of human vector population is still a challenge as the vector has a diurnal feeding and outdoor resting behavior. Environmental management is still the best practice to be adopted in many countries for Aedes aegypti control. The currently discovered dengue vaccine might be an immediate arsenal for the community immunization.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/64007",risUrl:"/chapter/ris/64007",signatures:"Eliningaya J. Kweka, Vito Baraka, Leah Mathias, Beda Mwang’onde,\nGermana Baraka, Lucile Lyaruu and Aneth M. Mahande",book:{id:"7107",title:"Dengue Fever",subtitle:"a Resilient Threat in the Face of Innovation",fullTitle:"Dengue Fever - a Resilient Threat in the Face of Innovation",slug:"dengue-fever-a-resilient-threat-in-the-face-of-innovation",publishedDate:"January 30th 2019",bookSignature:"Jorge Abelardo Falcón-Lezama, Miguel Betancourt-Cravioto and Roberto Tapia-Conyer",coverURL:"https://cdn.intechopen.com/books/images_new/7107.jpg",licenceType:"CC BY 3.0",editedByType:"Edited by",editors:[{id:"198399",title:"Dr.",name:"Jorge Abelardo",middleName:null,surname:"Falcón-Lezama",slug:"jorge-abelardo-falcon-lezama",fullName:"Jorge Abelardo Falcón-Lezama"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}},authors:[{id:"123576",title:"Prof.",name:"Eliningaya",middleName:null,surname:"Kweka",fullName:"Eliningaya Kweka",slug:"eliningaya-kweka",email:"pat.kweka@gmail.com",position:null,institution:null},{id:"225604",title:"MSc.",name:"Germana",middleName:null,surname:"Baraka",fullName:"Germana Baraka",slug:"germana-baraka",email:"germanathomas1@gmail.com",position:null,institution:null},{id:"251361",title:"Mr.",name:"Vito",middleName:null,surname:"Baraka",fullName:"Vito Baraka",slug:"vito-baraka",email:"vitobaraka@gmail.com",position:null,institution:null},{id:"251363",title:"Ms.",name:"Leah",middleName:null,surname:"Mathias",fullName:"Leah Mathias",slug:"leah-mathias",email:"leahmathiastz@gmail.com",position:null,institution:null},{id:"251365",title:"Ms.",name:"Lucile",middleName:null,surname:"Lyaruu",fullName:"Lucile Lyaruu",slug:"lucile-lyaruu",email:"lucillejustis@gmail.com",position:null,institution:null},{id:"251366",title:"Ms.",name:"Aneth",middleName:null,surname:"Mahande",fullName:"Aneth Mahande",slug:"aneth-mahande",email:"anethmmahande@gmail.com",position:null,institution:null},{id:"272719",title:"Dr.",name:"Beda",middleName:null,surname:"J. Mwang'onde",fullName:"Beda J. Mwang'onde",slug:"beda-j.-mwang'onde",email:"Bedajohnm@gmail.com",position:null,institution:null}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_1_2",title:"1.1. Aedes distribution",level:"2"},{id:"sec_2_2",title:"1.2. Ecology of Aedes mosquitoes",level:"2"},{id:"sec_2_3",title:"1.2.1. Aedes aegypti",level:"3"},{id:"sec_3_3",title:"1.2.2. Aedes albopictus",level:"3"},{id:"sec_4_3",title:"1.2.3. Aedes mosquitoes life cycle",level:"3"},{id:"sec_6_2",title:"1.3. Insecticide resistance in Aedes spp.",level:"2"},{id:"sec_7_2",title:"1.4. Mechanisms of insecticide resistance",level:"2"},{id:"sec_7_3",title:"1.4.1. Physiological resistance in Aedes spp.",level:"3"},{id:"sec_8_3",title:"1.4.2. Behavioral resistance in Aedes spp.",level:"3"},{id:"sec_11",title:"2. Disease transmission by Aedes aegypti",level:"1"},{id:"sec_12",title:"3. Seasonality and intensity of transmission",level:"1"},{id:"sec_13",title:"4. Control and surveillance",level:"1"},{id:"sec_13_2",title:"4.1. Community education",level:"2"},{id:"sec_14_2",title:"4.2. Larval mosquito control",level:"2"},{id:"sec_15_2",title:"4.3. Adult control of Aedes aegypti",level:"2"},{id:"sec_16_2",title:"4.4. Use of repellents",level:"2"},{id:"sec_17_2",title:"4.5. Surveillance",level:"2"},{id:"sec_19",title:"5. Discussion",level:"1"},{id:"sec_20",title:"6. Conclusion",level:"1"}],chapterReferences:[{id:"B1",body:'Harbach R. Mosquito Taxonomic Inventory. 2013. p. 2015\n'},{id:"B2",body:'Wilkerson RC, Linton Y-M, Fonseca DM, Schultz TR, Price DC, Strickman DA. Making mosquito taxonomy useful: A stable classification of tribe Aedini that balances utility with current knowledge of evolutionary relationships. PLoS One. 2015;10:e0133602\n'},{id:"B3",body:'Kraemer MU, Sinka ME, Duda KA, Mylne AQ, Shearer FM, Barker CM, et al. 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The American Journal of Tropical Medicine and Hygiene. 2010;83:277-284\n'},{id:"B34",body:'Brengues C, Hawkes NJ, Chandre F, McCarroll L, Duchon S, Guillet P, et al. Pyrethroid and DDT cross-resistance in Aedes aegypti is correlated with novel mutations in the voltage-gated sodium channel gene. Medical and Veterinary Entomology. 2003;17:87-94\n'},{id:"B35",body:'WHO. Test Procedures for Insecticide Resistance Monitoring in Malaria Vector Mosquitoes. Geneva, Swirtzeland: WHO; 2016\n'},{id:"B36",body:'Feyereisen R. Insect CYP genes and P450 enzymes. In: Insect Molecular Biology and Biochemistry. London: Elsevier, Academic Press; 2012. pp. 236-316\n'},{id:"B37",body:'Stevenson BJ, Pignatelli P, Nikou D, Paine MJ. Pinpointing P450s associated with pyrethroid metabolism in the dengue vector, Aedes aegypti: Developing new tools to combat insecticide resistance. 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A mutation in the voltage-gated sodium channel gene associated with pyrethroid resistance in Latin American Aedes aegypti. Insect Molecular Biology. 2007;16:785-798\n'},{id:"B42",body:'Kawada H, Higa Y, Komagata O, Kasai S, Tomita T, Yen NT, et al. Widespread distribution of a newly found point mutation in voltage-gated sodium channel in pyrethroid-resistant Aedes aegypti populations in Vietnam. PLoS Neglected Tropical Diseases. 2009;3:e527\n'},{id:"B43",body:'David J-P, Ismail HM, Chandor-Proust A, Paine MJI. Role of cytochrome P450s in insecticide resistance: Impact on the control of mosquito-borne diseases and use of insecticides on earth. Philosophical Transactions of the Royal Society, B: Biological Sciences. 2013;368\n'},{id:"B44",body:'Faucon F, Gaude T, Dusfour I, Navratil V, Corbel V, Juntarajumnong W, et al. In the hunt for genomic markers of metabolic resistance to pyrethroids in the mosquito Aedes aegypti: An integrated next-generation sequencing approach. PLoS Neglected Tropical Diseases. 2017;11:e0005526\n'},{id:"B45",body:'Hemingway J, Hawkes NJ, McCarroll L, Ranson H. The molecular basis of insecticide resistance in mosquitoes. Insect Biochemistry and Molecular Biology. 2004;34:653-665\n'},{id:"B46",body:'Lumjuan N, Rajatileka S, Changsom D, Wicheer J, Leelapat P, Prapanthadara L-A, et al. The role of the Aedes aegypti Epsilon glutathione transferases in conferring resistance to DDT and pyrethroid insecticides. Insect Biochemistry and Molecular Biology. 2011;41:203-209\n'},{id:"B47",body:'Hemingway J, Ranson H. Insecticide resistance in insect vectors of human disease. Annual Review of Entomology. 2000;45:371-391\n'},{id:"B48",body:'Malik A, Earhart K, Mohareb E, Saad M, Saeed M, Ageep A, et al. Dengue hemorrhagic fever outbreak in children in Port Sudan. Journal of Infection and Public Health. 2011;4:1-6\n'},{id:"B49",body:'Amarasinghe A, Kuritsky JN, Letson GW, Margolis HS. Dengue virus infection in Africa. Emerging Infectious Diseases. 2011;17:1349\n'},{id:"B50",body:'Gubler DJ. Dengue and dengue hemorrhagic fever. Clinical Microbiology Reviews. 1998;11:480-496\n'},{id:"B51",body:'Gubler DJ, Clark GG. Community-based integrated control of Aedes aegypti: A brief overview of current programs. The American Journal of Tropical Medicine and Hygiene. 1994;50:50-60\n'},{id:"B52",body:'Thongcharoen P, Jatanasen S. Epidemiology of dengue and dengue haemorrhagic fever. Monograph on Dengue/Dengue Haemorrhagic Fever. 1999:1-8\n'},{id:"B53",body:'Kroeger ALA, Ochoa M, Villegas E, Levy M, Alexander N. Effective control of dengue vectors with curtains and water container covers treated with insecticide in Mexico and Venezuela: Cluster randomised trials. BMJ. 2006;332:1247\n'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Eliningaya J. Kweka",address:"eliningaya.kweka@tpri.go.tz",affiliation:'
School of Medicine, Department of Parasitology and Entomology, Catholic University of Health and Allied Health Sciences, Tanzania
'},{corresp:null,contributorFullName:"Aneth M. Mahande",address:null,affiliation:'
Tropical Pesticides Research Institute, Mabogini Field Station, Tanzania
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The first-principle modeling approach combined with the frequency system identification has been adopted to obtain a high-fidelity dynamics model. It is inherently less stable and difficult to control. To accomplish the required practical flight tasks, the flying vehicle needs to work well even in windy conditions. Moreover, for flight control engineers, simple prescribed multi-loop controller structures are preferred. To handle the multiple problems, a structured velocity controller consisting of two feedback loops is developed, where inner loop provides stability augmentation and decoupling, and the outer loop guarantees desired velocity tracking performance. The simultaneous design of the two-loop controllers under multiple performance requirements in the usual H∞ metrics can be cast as a nonsmooth optimization program. To compensate for changes in plant dynamics across the flight envelope, a smooth and compact polynomial scheduling formula is implemented as a function of the forward flight speed. Both simulations and flight test results have been presented in this work to showcase the potential for the proposed robust nonlinear control system to optimize the performance of UAV, specifically unconventional vehicles.",signatures:"Yang Wang, Changle Xiang, Yue Ma and Bin Xu",authors:[{id:"181878",title:"Ph.D.",name:"Yang",surname:"Wang",fullName:"Yang Wang",slug:"yang-wang",email:"yangwangeducn@gmail.com"},{id:"186663",title:"Prof.",name:"Changle",surname:"Xiang",fullName:"Changle Xiang",slug:"changle-xiang",email:"xiangcl@bit.edu.cn"},{id:"186664",title:"Prof.",name:"Yue",surname:"Ma",fullName:"Yue Ma",slug:"yue-ma",email:"yuema.bit@gmail.com"},{id:"186665",title:"Dr.",name:"Bin",surname:"Xu",fullName:"Bin Xu",slug:"bin-xu",email:"bitxubin@bit.edu.cn"}],book:{title:"Autonomous Vehicle",slug:"autonomous-vehicle",productType:{id:"1",title:"Edited Volume"}}}],collaborators:[{id:"24557",title:"Prof.",name:"Shengbo",surname:"Li",slug:"shengbo-li",fullName:"Shengbo Li",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"176926",title:"Mr.",name:"Khuram",surname:"Shahzad",slug:"khuram-shahzad",fullName:"Khuram Shahzad",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/176926/images/4278_n.jpg",biography:"Mr. Khuram Shahzad is an enthusiastic, competent, reliable and fast-learning person with a broad and acute interest in computer science, programming, robotics and automation, mathematical modeling, artificial intelligence and machine learning. He has recently completed his Masters studies from National University of Sciences and Technology (NUST), Islamabad, Pakistan. During his Masters Research thesis, he has successfully completed research on “Safe Path Planning for a Continuum Robot for Surgical Applications”. During his Bachelor studies, he secured first position (Gold Medal) in Bachelor of Science (Computer Science) amongst all the candidates of his batch. Mr. Khuram Shahzad is currently employed in Business Analytics Pvt. Ltd. Islamabad, Pakistan as “Senior Software Engineer”. He is working with Business Analytics (BA) Pvt. Ltd. from September 2011. During his tenure at BA, he has worked on design and development of different range of projects, including commercial cloud mailing server (admin as well as client development), mobile apps, shopping cart, communication server and helpdesk. He is project lead for different ongoing projects in the organization. Because of his excellent job performance, he has been awarded with appreciation award (March 2015) from his employer (Business Analytics Pvt. Ltd. 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The Open Access model is applied to all of our publications and is designed to eliminate subscriptions and pay-per-view fees. This approach ensures free, immediate access to full text versions of your research.
As a gold Open Access publisher, an Open Access Publishing Fee is payable on acceptance following peer review of the manuscript. In return, we provide high quality publishing services and exclusive benefits for all contributors. IntechOpen is the trusted publishing partner of over 118,000 international scientists and researchers.
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The Open Access Publishing Fee (OAPF) is payable only after your full chapter, monograph or Compacts monograph is accepted for publication.
\\n\\n
OAPF Publishing Options
\\n\\n
\\n\\t
1,400 GBP Chapter - Edited Volume
\\n\\t
10,000 GBP Monograph - Long Form
\\n\\t
4,000 GBP Compacts Monograph - Short Form
\\n
\\n\\n
*These prices do not include Value-Added Tax (VAT). Residents of European Union countries need to add VAT based on the specific rate in their country of residence. Institutions and companies registered as VAT taxable entities in their own EU member state will not pay VAT as long as provision of the VAT registration number is made during the application process. This is made possible by the EU reverse charge method.
\\n\\n
Services included are:
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\\n\\t
An online manuscript tracking system to facilitate your work
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Assurance that your manuscript meets the highest publishing standards
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English language copyediting and proofreading, including the correction of grammatical, spelling, and other common errors
\\n\\t
XML Typesetting and pagination - web (PDF, HTML) and print files preparation
\\n\\t
Discoverability - electronic citation and linking via DOI
\\n\\t
Permanent and unrestricted online access to your work
What isn't covered by the Open Access Publishing Fee?
\\n\\n
If your manuscript:
\\n\\n
\\n\\t
Exceeds 20 pages (for chapters in Edited Volumes), an additional fee of 40 GBP per page will be required
\\n\\t
If a manuscript requires Heavy Editing or Language Polishing, this will incur additional fees.
\\n
\\n\\n
Your Author Service Manager will inform you of any items not covered by the OAPF and provide exact information regarding those additional costs before proceeding.
\\n\\n
Open Access Funding
\\n\\n
To explore funding opportunities and learn more about how you can finance your IntechOpen publication, go to our Open Access Funding page. IntechOpen offers expert assistance to all of its Authors. We can support you in approaching funding bodies and institutions in relation to publishing fees by providing information about compliance with the Open Access policies of your funder or institution. We can also assist with communicating the benefits of Open Access in order to support and strengthen your funding request and provide personal guidance through your application process. You can contact us at oapf@intechopen.com for further details or assistance.
\\n\\n
For Authors who are still unable to obtain funding from their institutions or research funding bodies for individual projects, IntechOpen does offer the possibility of applying for a Waiver to offset some or all processing feed. Details regarding our Waiver Policy can be found here.
\\n\\n
Added Value of Publishing with IntechOpen
\\n\\n
Choosing to publish with IntechOpen ensures the following benefits:
\\n\\n
\\n\\t
Indexing and listing across major repositories, see details ...
\\n\\t
Long-term archiving
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Visibility on the world's strongest OA platform
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Live Performance Metrics to track readership and the impact of your chapter
\\n\\t
Dissemination and Promotion
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\\n\\n
Benefits of Publishing with IntechOpen
\\n\\n
\\n\\t
Proven world leader in Open Access book publishing with over 10 years experience
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+4,800 OA books published
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Most competitive prices in the market
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Fully compliant with OA funding requirements
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Optimized processes, enabling publication between 8 and 12 months
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Personal support during every step of the publication process
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+146,150 citations in Web of Science databases
\\n\\t
Currently strongest OA platform with over 130 million downloads
As a gold Open Access publisher, an Open Access Publishing Fee is payable on acceptance following peer review of the manuscript. In return, we provide high quality publishing services and exclusive benefits for all contributors. IntechOpen is the trusted publishing partner of over 118,000 international scientists and researchers.
\n\n
The Open Access Publishing Fee (OAPF) is payable only after your full chapter, monograph or Compacts monograph is accepted for publication.
\n\n
OAPF Publishing Options
\n\n
\n\t
1,400 GBP Chapter - Edited Volume
\n\t
10,000 GBP Monograph - Long Form
\n\t
4,000 GBP Compacts Monograph - Short Form
\n
\n\n
*These prices do not include Value-Added Tax (VAT). Residents of European Union countries need to add VAT based on the specific rate in their country of residence. Institutions and companies registered as VAT taxable entities in their own EU member state will not pay VAT as long as provision of the VAT registration number is made during the application process. This is made possible by the EU reverse charge method.
\n\n
Services included are:
\n\n
\n\t
An online manuscript tracking system to facilitate your work
\n\t
Personal contact and support throughout the publishing process from your dedicated Author Service Manager
\n\t
Assurance that your manuscript meets the highest publishing standards
\n\t
English language copyediting and proofreading, including the correction of grammatical, spelling, and other common errors
\n\t
XML Typesetting and pagination - web (PDF, HTML) and print files preparation
\n\t
Discoverability - electronic citation and linking via DOI
\n\t
Permanent and unrestricted online access to your work
What isn't covered by the Open Access Publishing Fee?
\n\n
If your manuscript:
\n\n
\n\t
Exceeds 20 pages (for chapters in Edited Volumes), an additional fee of 40 GBP per page will be required
\n\t
If a manuscript requires Heavy Editing or Language Polishing, this will incur additional fees.
\n
\n\n
Your Author Service Manager will inform you of any items not covered by the OAPF and provide exact information regarding those additional costs before proceeding.
\n\n
Open Access Funding
\n\n
To explore funding opportunities and learn more about how you can finance your IntechOpen publication, go to our Open Access Funding page. IntechOpen offers expert assistance to all of its Authors. We can support you in approaching funding bodies and institutions in relation to publishing fees by providing information about compliance with the Open Access policies of your funder or institution. We can also assist with communicating the benefits of Open Access in order to support and strengthen your funding request and provide personal guidance through your application process. You can contact us at oapf@intechopen.com for further details or assistance.
\n\n
For Authors who are still unable to obtain funding from their institutions or research funding bodies for individual projects, IntechOpen does offer the possibility of applying for a Waiver to offset some or all processing feed. Details regarding our Waiver Policy can be found here.
\n\n
Added Value of Publishing with IntechOpen
\n\n
Choosing to publish with IntechOpen ensures the following benefits:
\n\n
\n\t
Indexing and listing across major repositories, see details ...
\n\t
Long-term archiving
\n\t
Visibility on the world's strongest OA platform
\n\t
Live Performance Metrics to track readership and the impact of your chapter
\n\t
Dissemination and Promotion
\n
\n\n
Benefits of Publishing with IntechOpen
\n\n
\n\t
Proven world leader in Open Access book publishing with over 10 years experience
\n\t
+4,800 OA books published
\n\t
Most competitive prices in the market
\n\t
Fully compliant with OA funding requirements
\n\t
Optimized processes, enabling publication between 8 and 12 months
\n\t
Personal support during every step of the publication process
\n\t
+146,150 citations in Web of Science databases
\n\t
Currently strongest OA platform with over 130 million downloads
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