The electroplating parameters and the properties of the deposit
\r\n\tThere are different types of multiple pregnancies: fraternal twins, identical twins, triplets, and higher-order multiples. Symptoms of multiple pregnancies are larger uterus than expected for the date in pregnancy, increased morning sickness, increased appetite, and excessive weight gain. In this book, we will examine the clinical aspects of multiple pregnancies and management. Also, we will examine the management of cases of twins including antenatal care, delivery, and postpartum.
",isbn:"978-1-80356-198-1",printIsbn:"978-1-80356-197-4",pdfIsbn:"978-1-80356-199-8",doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!0,isSalesforceBook:!1,hash:"70396c6f5f2928c422c1eaf6d33c6269",bookSignature:"Prof. Hassan S Abduljabbar",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/11732.jpg",keywords:"Multiple Pregnancies, Twins, Physiology, Incidence, Epidemiology, Demographics, Predisposing Factors, Prenatal Diagnosis, Zygosity, Complications, Management of Birth, Antenatal Care",numberOfDownloads:null,numberOfWosCitations:0,numberOfCrossrefCitations:null,numberOfDimensionsCitations:null,numberOfTotalCitations:null,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"February 4th 2022",dateEndSecondStepPublish:"May 4th 2022",dateEndThirdStepPublish:"July 3rd 2022",dateEndFourthStepPublish:"September 21st 2022",dateEndFifthStepPublish:"November 20th 2022",remainingDaysToSecondStep:"14 days",secondStepPassed:!0,currentStepOfPublishingProcess:3,editedByType:null,kuFlag:!1,biosketch:"Professor of Obstetrics and Gynecology at King Abdulaziz University, Saudi Arabia, consultant, clinician, researcher, editor, and referee of many international scientific medical journals. Dr. Abduljabbar is president of the Saudi Society of Obstetrics and Gynecology and the president of the Federation of Arab Gynecology Obstetrics Societies. He has published more than seventy-five scientific articles and edited several books.",coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"68175",title:"Prof.",name:"Hassan",middleName:"S",surname:"Abduljabbar",slug:"hassan-abduljabbar",fullName:"Hassan Abduljabbar",profilePictureURL:"https://mts.intechopen.com/storage/users/68175/images/system/68175.png",biography:"Hassan S. Abduljabbar, MD, FRCSC, American Board Diplomate, is a professor at the College of Medicine, King Abdulaziz\nUniversity, Saudi Arabia. He is also the president of the Saudi Society of Obstetrics and Gynecology and the Federation of Arab\nGynecology Obstetric Societies (FAGOS). He is a referee for\nmany international scientific journals. 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The particles are trapped in the coating during deposition. Composite coatings are composed of an electrodeposited metal matrix and dispersed solid particles. The metal powders, metal alloy powders and metal oxide powders of Al, Co, Cu, In, Mg, Ni, Si, Sn, V, Zn and nitrides of Al, B, Si and C (graphite or diamond) and carbides of B, Bi, Si, W and MoS2 and organic materials such as polytetrafluoroethylene (PTFE) and polymer spheres are used as the particles [40].
The main application areas of the composite deposition are electronic, biomedical, telecommunication, automotive, space and consumer applications where high strength, equiaxed micro-components are required.
The electroplating parameters must be controlled during electrodeposition since they have significant effect on the coating properties, deposition efficiency and hydrogen evolution. Besides the current density, pH of the bath, bath temperature that places among the main electroplating parameters [36], there are more parameters including the type, the size, the concentration of the particles [37] and the addition of surfactants and their types [20]. The main properties of the coating can be listed as the corrosion resistance in TiO2 – nickel coating [38], tribological properties in MoS2 – nickel coating [4], mechanical properties in TiO2 – nickel coating [38], internal stress in MoS2 – nickel coating [9], texture in chromium – carbon deposit [39], particle content of the deposit [24] are severely affected by the electroplating parameters.
The aim of this chapter is to emphasize on the electroplating parameters, their effects and interaction effect on the coating properties. Particle incorporation in the deposit is an important property that must be analysed since composite electroplating aroused in order to improve the matrix properties with particle addition. Another important property is the internal stress that must kept at minimum levels not to disrupt the deposit.
Electrodeposition process consists of two steps that are nucleation, growth mechanisms and thickening of the primary layer. The nucleation is enhanced by high current density unlike the growth process [1]. Thus, smaller grain sizes are observed at higher current densities due to the increase in the nucleation rate [2]. On the other hand, high current density increases pH in the vicinity of the electrode during the reduction process that creates a competition between metal deposition and hydrogen gas [6] [3]. Hydrogen evolution contributes to the internal stress in the deposit [4][5]. Therefore, it is important to figure out the current density values at which hydrogen evolution does not occur together with metal deposition [4].
The current density also affects the particle content in the deposit. The particle (WC) content of the coating increases linearly with an increase in the current density from 0.1 to 0.5 A/dm2 regardless of the particle diameters [7]. On the other hand, Kuo [6] claims that the particle (MoS2) content in the deposit decreases when the current density increases from 4 to 8 A/dm2. According to the results of Hu and Bai [41], increasing current density generally increased the particle content but the effect of current density depends on pH. Figure 1 shows that when the current density (B) is increased, the particle content is increasedwhen pH (C) is 1 and decreased when pH is 5 [41].
The effects and interaction effects of parameters on the atomic percent of phosphorus in the deposit. A: main effect of temperature, B: main effect of current density, BxC: current density – pH interaction effect, AxB: temperature – current density interaction effect [
Stress behaviour for sulfamate nickel electrolytes with three different chemistries [
Another effect of increasing current density from 0.15 to 5 A/dm2 is increasing the internal stress [8] since the stress stems from the residual stresses. However significant effect cannot be detected when the current density increased from 1.2 to 4.8 A/dm2 [9] because the effect of current density on the internal stress depends on the electroplating solution composition. Figure 2 shows the effect of changing only one component amount of the solution on the current density dependence of the internal stress [42]. Moreover the effect of the current density depends on pH.
The effect of pH on particle incorporation is dependent on the nature of the particles. For instance, when MoS2 particles are used as the incorporate particles, the effect of pH value on the particle content in deposit was insignificant. Nevertheless it can be concluded that increasing pH decreased the amount of particles present in the coating [6]. However, incorporated particles are significantly decreased when pH is below 2 in Al2O3-Ni coating pair [11]. In addition, decreasing pH is preferred to manage the internal stress. Low pH values, less than 5, are selected to obtain acceptable stress levels [10]. Increase in pH of the solution may lead to discharge of hydroxyl ions instead of nickel dissolution and oxygen evolution [10] resulted in high internal stresses. It is also concluded that the internal stress was increased when the pH was increased from 2 to 4 in MoS2 – Ni system [12].
The effect of the temperature on the particle content in the deposit depends on the type of the particle. There is a small increase in the particle (MoS2) content in coating with increasing the temperature 30, 40, 50 respectively [6]. It was mentioned that the temperature has an insignificant effect. For instance, no effect of temperature was detected in BaCr2O4-Ni [13] and Al2O3-Ni [14] coating couples. Because, the applied voltage is the main parameter that directly affects the activity of the reaction. In addition, Ni deposits more efficiently with increase in temperature. On the other hand, the influence of the temperature was reported as positive up to certain point. After that point, the amount of particles decreases with increasing the temperature.
General trend on the effect of the bath temperature on the internal stress is positive, meaning that increasing temperature decreases the internal stress. On the other hand, according to some of the studies the stress is more influenced by the current density regardless of the temperature [10].
Another advantage of the high temperature is the polarization effect. It is known that concentration polarization is the component of the polarization which is due to the change in the electrolyte concentration that stems from the current flow through the electrode – solution interface. So, the electrochemical cell potential difference deviates from its equilibrium value. Concentration polarization is decreased by increasing temperature because diffusion layer thickness gets smaller and ionic diffusion increases.
On the other hand, high temperature values increases energy consumption and supply heat for bath evaporation. Furthermore, thermal stresses will arrive at high processing temperatures and can become a serious problem especially when the coating and substrate have different thermal expansion coefficients. Therefore, an optimum plating temperature must be preferred to satisfy energy consumption and the coating quality.
It is a common fact that if the amount of the particles in the solution is increased, the particle content in the deposit will also increase up to a certain point. However the type of the particle also acts as an important parameter. Both of the conductive and non-conductive particles have their own advantages against each other. Because conducting particles (molybdenum disulfide, chromium carbide, zirconium diboride, graphite) attracted to the cathode then act as depositing sites which resulted in dendritic growth [15]. Despite the advantage of easily attraction of conducting particles to the cathode, selective deposition on the conducting sites led to increased surface roughness.
Schematic view of the composite coating including a) conducting and b)non-conducting particles
On the other hand, non-conductive particles end up with smoother deposit surfaces with low porosity [15].
Mechanical properties can be changed by the type of the incorporated particles. For instance, PTFE in Ni matrix increases wear resistance [16], MoS2 incorporated with Ni decreased the coefficient of friction [9], Al2O3 and SiC dispersed in Cu increased the microhardness of the coatings [17]. Moreover, wear and corrosion resistance can be improved by the addition of silicon carbide nano-particles [18] [19].
Small particle sizes can be agitated easily and led to an increase in the particle concentration in the deposit during composite electroplating. For example, the amount of particles is increased by decreasing the particle size in Ni/SiC system [21]. Furthermore, the effect of particles in decreasing the friction coefficient is more effective when the particle size is decreased in Ni-MoS2 system [9].
Generally, increasing the particle concentration in the bath increases the weight percentage of particles in the deposit up to a certain point [9] [22] [23]. That point can be thought like the saturation point. There is a rapid increase in the particle amount in the low particle concentration regions whereas a slight increase occurs in the high concentration regions. The collisions between particles and cathode determine the codeposition of the particles and they are diminished in the high MoS2 concentration region resulting in slight increase or decreased particle amount in the growing metal deposit [6].
Coating performance can be developed by the addition of the surfactants like (cetyltrimethylammonium bromide (CTAB), sodyumdodecyl sulfate (SDS), and saccharine [20]. The advantage of adding surfactants is their dispersing effect of particles. Thus the property of the particles will be uniform on the surface. Surfactants adsorb on the particles and favor the distribution of the particles [24].
Surfactants are indispensable especially for the hydrophobic particles (fluorographite, MoS2) to be dispersed in the electroplating solution. Surfactants like sodium lauryl sulphate enhanced the electrostatic adsorption of suspended particles on cathode surface by increasing their positive charges [25]. Similarly, azobenzene (AZTAB) promoted particle deposition into the nickel matrix by their more positive reduction potential than that of nickel [26]. Another surfactant, cetyl trimethyl ammonium bromide (CTAB) has an advantage of increasing the volume percentage of SiC in the deposit besides homogeneous and non-agglomerated distribution of particles in SiC-nickel composite coatings [27].
Further advantage of the surfactants is suppressing the hydrogen evolution reaction. For example, saccharin which is an anionic surfactant is an effective way to overcome the hydrogen evolution problem [34].
The surfactants can be grouped under two main headings which are anionic and cationic surfactants according to their charges. Cationic surfactants increased the particle incorporation in the coating [28] [29] [30] [31]. Anionic surfactants may have positive or negative effect on the codeposition efficiency of the particles depending on the particle type and bath solution. For instance, SDS which is an anionic surfactant did not affect the codepositon of particles [31]. On the other hand, cationic surfactants have the advantage of adsorbing on the particles that have negative surface charge [29]. Therefore, a net positive charge was formed by the adsorption of cationic surfactants that inhibited the formation of particle clusters and led to more stable particle suspension in the bath. Moreover, this positive charge improved the tendency of particles to move towards cathode and increased the amount of particles in the deposit [29] [30]. For instance, addition of cationic surfactant benzyl ammonium salts (BAS), increased the amount of MoS2 codeposition [32]. In addition, BAS adsorbed on MoS2 particles decreased the conductivity of the particles and resulted in homogeneous deposition of nickel and MoS2 particles [32] [15]. On the other hand, it was stated that anionic surfactants in the electrolyte can give particles a negative charge and make them to move towards the substrate [35].
The disadvantage of the surfactants may occur if there are unabsorbed free surfactants, because they could lead to stress development and brittleness in the deposit [33]. Since the amount of incorporated surfactant is generally very small, their undesirable effects may be ignored [33]. However, increasing surfactant (CTAB) amount caused an increase in the internal stress due to the high possibilty of embedded CTAB in the nickel matrix [32].
2n-1 fractional factorial design is a statistical design that can be used to identify the effects of n electrodeposition variables on the coating properties with the reasonable number of experiments. In addition, the interaction effects of the parameters can be analyzed by the help of fractional factorial. The property of the coating in other words the response value in the program is generally taken as the amount of particles in the deposit during composite coatings. For instance, in the study of Hu and Bai [41], phosphorus content in the deposit was taken as the response value and temperature, current density pH, NaH2PO2 H2O concentration of the solution and agitation rate were taken as the electroplating parameters. Another most commonly used response value during composite coating is internal stress. Electroplating parameters were MoS2 particle concentration, temperature, pH, current density and coating thickness where the response value is the internal stress in the study of Saraloglu Guler et. al. [9]. Other response values can be listed as friction coefficient, corrosion resistance, wear resistance, hardness which are the properties obtained by particle addition so increased amount of particle content in the deposit will have a positive effect on these values. The hydrogen evolution reaction must also be considered during this selection. The effects of the electroplating parameters on hydrogen evolution reaction can be studied before the composite deposition in order to determine the current density range where H2 is not simultaneously discharged with Ni plating [12].
Fixed limit values that are said to be low (-1) and high (1) levels are selected for the electroplating parameters in fractional factorial design. Table 1 shows the parameters and their low and high fixed limit values for levels of fractional factorial design.
\n\t\t\t\t | \n\t\t\t\n\t\t\t\t | \n\t\t
Current density | \n\t\t\tThe coefficient of friction | \n\t\t
pH of the bath | \n\t\t\tWear resistance | \n\t\t
Bath temperature | \n\t\t\tCorrosion resistance | \n\t\t
Particle concentration of the bath | \n\t\t\tMechanical Properties | \n\t\t
Particle size | \n\t\t\tInternal Stress | \n\t\t
Particle type | \n\t\t\tTexture formation | \n\t\t
Surfactant addition/ type | \n\t\t\tParticle content of the deposit | \n\t\t
\n\t\t\t | Grain size | \n\t\t
\n\t\t\t | Electrodeposition/ particle uniformity | \n\t\t
The electroplating parameters and the properties of the deposit
Photosynthetic eukaryotic organisms harbor a chloroplast genome (also called ‘plastome’) within their cells. This genome derives from the endosymbiosis of a prokaryotic organism, which was then gradually converted into the chloroplast. With the increased number of sequences within publicly available databases and the emergence of very sophisticated phylogenetic and phylogenomic analyses, we can infer much more precisely the origin of this primary endosymbiotic event. In addition, these comparative analyses allow for investigation of plastome evolutionary dynamics in the different plant lineages and the extent of nuclear influence over the chloroplast genome. Overall, plant plastomes harbor a very low gene content compared to their prokaryotic ancestor, which appears to result from either gene loss due to redundant functions in both chloroplast and nuclear genomes or functional transfer and relocation of chloroplast genes into the nucleus. The relocation of thousands of chloroplast genes from the chloroplast to the nucleus was rendered possible due to the massive transfer of DNA from the chloroplast to the nucleus. However, chloroplast genes that have been integrated into the nucleus are not immediately functional and have to adapt to their new eukaryotic environment by acquiring various regulatory elements (i.e., promoter, polyadenylation signal, and target peptide). Despite most of these functional transfers occurred soon after the endosymbiotic event, some clever real-time experiments (using a selectable marker) have allowed for understanding how easily and by which molecular mechanisms DNA is transferred from the chloroplast to the nucleus. Such experiments have also permitted the study of the subsequent evolution of chloroplast DNA in the nuclear genome, and how a chloroplast gene becomes functional in the nucleus.
Photosynthetic eukaryotic cells arose through the engulfment of a cyanobacterium that was then converted into the chloroplast, enabling plants to use sunlight to fix carbon. This major functional innovation allowed for eukaryotes to transition from heterotrophy to autotrophy. This primary endosymbiotic event is at the origin of the astonishing biodiversity visible today in plants, including the Glaucophyta, Rhodophyta, and Viridiplantae lineages (Figure 1). With the advent of next-generation sequencing technologies, the number of fully sequenced plastomes has hugely expanded, providing insight into chloroplast evolution in the different plant lineages. In this part, we will present our current knowledge on chloroplast origin and what has been unraveled on the chloroplast genome evolution, regarding genome size, gene content, structure, and mutation rate.
Phylogenetic relationships of the different plant lineages formed after the primary endosymbiosis of a cyanobacterium by an ancestor of the Archaeplastida. The number of available genomes on GenBank is indicated under the image. For simplicity, “Mosses, Marchantyophytes and Bryophytes” on one side, as well as “Ferns and Lycopodiophyta” on the other side, were grouped together in the tree. Pictures copyright to L. Brient, M.T. Misset, R. Delourme, and J. Keller.
The first hypothesis of the endosymbiotic origin of chloroplasts is commonly credited to Russian botanist K. Mereschkowsky, who observed similarities between cyanobacteria and chloroplasts of plants and algae [1]. This hypothesis was then reaffirmed by Margulis in the 1970s. The origin of this primary endosymbiosis event is still debated. While fossil-based phylogeny estimated the origin of chloroplasts to be around 1.4–1.7 billion years ago [2], gene-based approaches dated it around 0.9 billion years ago [3]. Different phylogenetic analyses aimed at determining the cyanobacterial lineage from which the chloroplast was derived and revealed that chloroplasts were closely related to the nitrogen-fixing cyanobacteria Chroococcales,
It is now widely accepted that this primary endosymbiotic event has a single origin [6, 7, 8]; however, it is still unclear how long it took for the conversion of the bacterial endosymbiont into a fully integrated organelle. This transition from endosymbiont to organelle surely involved many steps. The first steps corresponded to the loss of the bacterial wall and the early acquisition by the endosymbiont of a transport system to transfer proteins and metabolites from the cytosol to the chloroplast. This latter step is constituted by two protein complexes: translocon of the outer (TOC) membranes of the chloroplast and translocon of the inner (TIC) membranes of the chloroplast [9, 10, 11]. The TIC/TOC complexes allow for transportation of the pre-proteins (proteins with a cleavable chloroplast target peptide) from the cytosol, where they are synthetized, to the chloroplast, where the target peptide is cleaved (reviewed in [11]). The presence of the same protein import apparatus in the different Archaeplastida lineages is the best evidence of the single origin of chloroplasts. Finally, the transition also necessitated the gradual functional transfer of endosymbiont genes to the nucleus [12], leading to the massive reduction of plastome size and gene content.
Most of our current knowledge of the conversion from endosymbiont to organelle has been obtained by comparing contemporary Archaeplastida organelles with their closest bacterial relatives. During the last few years, advances in high-throughput sequencing and bioinformatic methods greatly facilitated the assembly, analysis, and publication of complete plastomes. To date, more than 2300 plastomes are fully assembled and deposited in the GenBank database. This number of plastomes actually doubled in the last 2 years. However, the number of sequenced plastomes varies greatly between the different Archaeplastida lineages. Indeed, almost 80% of them belong to Angiosperms. Thus, there is an important inequality in the sequencing effort. The poor level of plastome sequencing in plant lineages outside of the Angiosperms needs to be improved to fully understand chloroplast genome evolution in plants. Some efforts to fill this gap have been performed in the last 2–5 years, but they are still insufficient. In the Glaucophyta, only one chloroplast genome is available (NC_001675), and another is sequenced but not yet published (Lang et al., unpublished). In contrast, the sequencing of Rhodophyta and Chlorophyta (green algae
Cumulative numbers of full chloroplast genomes deposited in GenBank for (A) Rhodophyta, (B) Chlorophyta, and (C) Streptophyta.
As mentioned previously, the conversion of the cyanobacterial endosymbiont into a chloroplast necessitated the functional transfer or replacement of most cyanobacterial genes into the nucleus. Compared to the thousands of genes (at least 2000) thought to have been once present in the cyanobacterial genome, Archaeplastida plastomes encode a maximum of around 250 genes [13, 14]. This observation indicates that most genes (includes protein coding and structural RNAs) present in the cyanobacterial ancestor have been functionally transferred relatively soon after the endosymbiotic event. Despite gene content among modern chloroplast genomes being relatively well conserved, there are important variations. Thus, Rhodophyta have the highest number of genes (237 in average; minimum 207; up to 266 in
Plastome numbers and characteristics (average size, number of proteins, and structural RNAs) among the Archaeplastida. The minimum and maximum genome sizes are indicated in italic.
These variations in gene content revealed the divergent evolution of plastomes in the different lineages. As an example, Rhodophyta gene content is characterized by the complete absence of the NADPH dehydrogenase complex [15]. Conversely, some genes are Rhodophyta-specific or rare in other Archaeplastida such as RNase P RNA, tmRNA, or signal recognition particle RNA [16, 17, 18]. Rhodophyta chloroplasts generally have a large genome size (see later) characterized by a high number of genes and other features such as the presence of bacteria-like operons, suggesting that Rhodophyta plastomes are phylogenetically closest to the ancestral cyanobacteria genome than any other algae [15]. Gene content variations are also well documented in the Angiosperm family in which multiple independent gene losses have been found such as
Among plants, chloroplast genomes range from less than 100 kb to more than 1 Gb, again excluding the non-chlorophyll species that exhibit significantly smaller chloroplast genomes (Table 1). The largest chloroplast genome ever sequenced has very recently been found in the red algae
Several factors can explain the important size variations found among the Archaeplastida. In the case of the red algae
Among plants, most plastomes seem to exhibit a conserved quadripartite structure, with a large and small single copy separated by two inverted repeats (Palmer 1983). However, multiple rearrangements occurred in diverse lineages, which modified this conserved structure. One of the most striking examples is the loss of one IR that occurred multiple times in the different chloroplast-bearing lineages, such as in the Fabaceae and the Geraniaceae [30, 33, 34]. This has also been reported for different Gymnosperms species such as
Chloroplast genome structure and gene order are also highly affected by inversions. Many inversions have been described in the literature, especially in legumes, with, for instance, fragments of 50 kb in the Papilionoideae [38], 36 kb in the Genistoids [39]; 29 kb in Sophoreae [40] or 7 kb in
Chloroplast genomes are known to be highly conserved, with relatively low rates of mutations, especially when compared to the plant nuclear genome. Indeed, the chloroplast genome evolves on average 10 times slower than the nuclear genome [45], with about 1 or less mutation/kb/million years [46] compared with approximately 7 mutations/kb/million years for the nuclear genome [47]. However, there are some exceptions, especially in three Angiosperm families (i.e., Fabaceae, Campanulaceae, and Geraniaceae) that are known to have accelerated evolutionary rates of their plastomes along with multiple structural rearrangements and size variations [19, 28, 30, 42, 44, 48, 49]. For example, the
To sum up this first section on the origin and evolution of plant plastomes originating from the primary endosymbiosis event, the recent sequencing and bioinformatics progress significantly increased the number of chloroplast genomes available for the scientific community. These advances have greatly improved our knowledge about the evolutionary dynamics of plastomes. Despite the diversity of organisms that harbor chloroplasts, plastomes in general seem to be relatively well conserved among the Archaeplastida (in terms of structure, size, and gene content); however, multiple independent alterations of these features have been observed in the different lineages. In addition, a few plant families (or group of species) seem to present an atypical evolution of the chloroplast genome. It is certain that the continuous effort to sequence much more plastomes (especially in the Glaucophyta and Rhodophyta) will allow the identification of new examples of such atypical evolution and will permit a better understanding of what are the causes and the molecular mechanisms involved in limiting or increasing plastome evolution.
Since the endosymbiotic event, the host genome (nuclear) has acquired most of the cyanobacterial genes, leading to the gradual loss of autonomy of the endosymbiont and the reduction of its genome. In this part, we will present our current knowledge on the mechanisms as well as the numerous cases of chloroplast DNA transfers to the nucleus and where it is now integrated in the nuclear genome. We will then detail the subsequent evolution and adaptation processes of the chloroplast genome that took place in its new eukaryotic environment. We will also discuss which factors can influence relocation of a chloroplast gene to the nucleus, and how a chloroplast gene transferred to the nucleus may become functional. Finally, we will discuss the important role that transfer of chloroplast DNA to the nucleus plays in the process of diversifying the plant nuclear gene content.
Much earlier than the complete sequencing and assembly of the first chloroplast genome (
Some of the chloroplast DNA fragments that were experimentally shown to insert in the nuclear genome were characterized [55, 60] and were often large in size (usually greater than 10 kb in length). Considering the massive transfer of chloroplast DNA to the nucleus, one would expect that some of these
Following endosymbiosis, the symbiont to organelle transition involved many steps. This includes the loss of the bacterial cell wall, the acquisition of a protein machinery that transfers nuclear-encoded proteins from the cytosol to the chloroplast (also known as the TIC and TOC complexes [75, 76]), and finally, the functional relocation of most chloroplast genes to the nucleus. As detailed below, a chloroplast gene may be replaced either only after its functional transfer to the nucleus, or directly substituted by a gene of a mitochondrial or eukaryotic origin.
Since the endosymbiosis event, thousands of genes have relocated within the nuclear genome. Indeed, cyanobacterial genomes encode a minimum of 2000 proteins, whereas current plant plastomes encode only 80–200 proteins, although 800 to more than 2000 proteins have been found in some algae and plant chloroplasts [77], respectively. Apart from some genes that presented redundant functions in both chloroplast and nuclear genomes, most chloroplast genes have been functionally relocated to the nucleus with their proteins targeted back to the organelle. Thus, the spectrum of proteins required for function and biogenesis of the cytoplasmic organelle did not greatly evolve since its creation.
The current plastome of most plants encodes a maximum of 200 proteins [78] whereas more than 2000 proteins in the chloroplast, suggesting the functional gene transfer and relocation of most chloroplast genes to the nucleus. As chloroplast genes are of prokaryote origin, they are not readily functional in the nuclear genome. To function in this novel environment, a chloroplast gene has to acquire or hijack nuclear gene regulatory elements (eukaryote promoter and terminator), as well as a transit peptide to target the protein back to the chloroplast [60, 79]. However, the acquisition of all these nuclear elements does not have to take place right after the transfer of the chloroplast gene to the nucleus, as they can retain their open reading frames for several million years [71]. In addition, some chloroplast genes can be relatively easily functional as a few chloroplast promoters (i.e.,
To date, the number of chloroplast-encoded proteins (about 80) is relatively well conserved among flowering plants. However, a few chloroplast genes have been independently lost in various plant lineages [19], allowing to understand how they became functional. Such chloroplast gene losses were most particularly observed in the Fabaceae, for which the plastome has been extensively reorganized and contains localized accelerated mutation rates [19]. Some of these genes, such as
The functional replacement of a chloroplast gene does not necessarily necessitate its functional transfer from the chloroplast to the nucleus. In the case of the chloroplast RPS16 protein, the chloroplast
Another evolutionary mechanism enabling the functional replacement of a chloroplast gene may occur
The continuous deluge of organellar DNA to the nucleus has facilitated the functional transfer of almost all chloroplast genes to the nucleus, reducing extensively the plastome size. Additionally, this organellar DNA was not only used to replace organellar genes but also enabled diversifying the plant nuclear gene content [77].
Chloroplast gene sequences transferred to the nucleus may present different fates. As presented in the two previous sections: (i) they may remain non-functional, decay, and ultimately be lost; (ii) they may acquire all the necessary elements to conserve the same function and have the protein targeted back to the chloroplast; or (iii) they may acquire new subcellular locations and functions. As mentioned earlier, Martin
The conversion of the cyanobacterial endosymbiont into the chloroplast partly results from the gradual transfer of hundreds to thousands of endosymbiont genes to the nuclear host. Across all lineages, more than 90% of the plant chloroplast proteins are now encoded in the nucleus. Within the few chloroplast-encoded proteins, about 40% of them are involved in chloroplast protein complexes that are made up of proteins encoded in both the chloroplast and the nucleus. These complexes exhibit important functions that are vital for the plant, such as photosystems I and II. One can only wonder how the stoichiometry between those two compartments is maintained. Indeed, one cell might contain hundreds to thousands of chloroplast copies compared to only one copy in the nucleus. Furthermore, chloroplast inheritance is often maternal, whereas nuclear bi-parental inheritance occurs in angiosperms during sexual reproduction. Therefore, coevolving interactions between cytoplasmic and nuclear genomes have been necessary and have resulted in significant coadaptation processes. When these fine-tuned coevolutionary interactions are disrupted, after intra-interspecific hybridization and/or genome doubling, for instance, incompatibilities and deleterious phenotypes can be observed. These evolutionary processes will be discussed in the light of previous work on synthetic and natural hybrids, as well as in polyploid species.
Several evolutionary scenarios can explain coadaptation between chloroplast and nuclear genomes after intraspecific hybridization. First, cytoplasmic genomes lack sexual reproduction and are more susceptible to fix and accumulate deleterious mutations by genetic drift [93]. Only positive selection for compensatory nuclear alleles will allow for regaining of optimal organelle function [94]. This mechanism of
Second, some mutations in the organelles could also be adaptive in specific environments and fixed in the population by natural selection. Subsequently, coadaptation process may favor specific nuclear variants to preserve intergenomic interactions. This mechanism is called
As mentioned above, the examples for intergenomic coadaptation and incompatibilities are scarce, and we are still very far from unraveling the molecular processes underlying such interactions. Applications of genome-wide studies in association with high-throughput sequencing would greatly improve our understanding of cytonuclear coevolution.
As shown above, cytonuclear interactions are extremely fine-tuned coevolved molecular processes that are still largely understudied. However, in recent years, efforts have been made, especially in neo-polyploid plant species (natural and resynthesized) to better apprehend the consequences of whole genome duplication (WGD) and interspecific hybridization on cytonuclear interactions and stability. In this last section, we will review our knowledge on such systems and elaborate on the many future issues to address.
Although completely overlooked, it is astonishing to envision the numerous and drastic consequences of a WGD event on copy number variation and stoichiometry on those cytonuclear complexes. Impacts of WGD on genomic structure and functional changes have been extensively studied in a large variety of plant systems. Genome redundancy can lead to changes in epigenetic patterns (including transposable element dynamics), altered gene expression (changes in global gene expression but also possible biased contribution of redundant copies), and fractionation processes (gene loss, homologous and non-homologous exchanges). However, to date, very few studies have investigated how the duplication of nuclear genes would affect the assembly dynamics of the multi-subunit cytonuclear complexes [103]. Different hypotheses predict the fate of nuclear and cytoplasmic genes implicated in cytonuclear complexes. They are based on the prediction that selection will favor compensatory mechanisms to maintain coordinated expression between cytoplasmic and nuclear genes leading
Only a handful of studies have looked at the consequences of WGD on a longer time scale, in that case, occurrences of subfunctionalization and pseudogenization of duplicated copies are to be expected. Coate et al. [109] stated that there might be a considerable influence of cytonuclear complex sensitivity to gene dosage imbalance and thus the need to return to single copy status or stay in duplicates. More specifically, Coate et al. [109] demonstrated that in
All of these processes could be enhanced through allopolyploidy, where divergent parental species first hybridized before genome doubling. In that case, the nuclear genome is redundant and a mixture of two, more or less, divergent parental genomes, whereas the organelles have (usually) a uniparental origin. Therefore, as chloroplast inheritance is usually maternal, selection should favor maintenance of maternal nuclear copies over the paternally inherited homoeolog as to preserve pre-existing coadaptive cytonuclear interactions. In allopolyploids, different scenarios leading to pseudogenization of paternal copies can be envisioned and were tested in a limited set of genes and species. The first scenario involves downregulation and relaxed selection of the paternally inherited homoeolog. An alternative scenario involves preferential gene conversion to the maternal homoeolog resulting in the loss of the paternal-like copy. It is important to note that both scenarios are not exclusive but could be part of a dynamic and gradual process, with first overexpression of the maternal copies leading to paternal homoeolog pseudogenization and maternally biased gene conversion. These hypotheses have only been tested in the Rubisco nuclear-encoded gene
These few studies already highlight the complexity of the different model systems that can be highly influenced by various evolutionary processes such as pre-existing coadaptive mechanisms, natural selection, and divergence between parental individuals (different populations to different species). As all Angiosperms have experienced at least one round of genome duplication and most of them multiple WGDs (Triticum and Brassica), paleopolyploid species are perfect candidates to elucidate the long-term impact of diploidization and biased genome fractionation on rates of asymmetric gene loss and pseudogenization. Additionally, it seems essential to integrate plant families that have contrasted rate of chloroplastic evolution (such as in Geraniaceae, Campanulaceae, and Fabaceae) and paternally inherited chloroplast genomes (such as in Actinidia, Medicago, and most Conifers). Finally, life history features such as reproductive strategy (perennial vs. annual), mating system (selfer vs. outcrosser), population level dynamics, and effective population size will also impact fixation rate of mutations.
We would like to thank the European Union Seventh Framework Program (FP7-CIG-2013-2017; Grant No. 333709 to Mathieu Rousseau-Gueutin) and an Agreenskills Plus fellowship to Julie Ferreira de Carvalho. We would also like to thank Dr. Christina Richards (Department of Integrative Biology, University of South Florida) for carefully and critical reading of the manuscript.
No conflict of interest.
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With data from the quarterly panel of the National Occupation and Employment Survey (NOES) of 2014–2016, we analyzed the mobility between eight categories: four of formal employment (non‐manual high‐skilled, non‐manual semi‐skilled, manual skilled manual and manual low‐skilled), two of informal employment (non‐manual and manual), unemployed and not in labor force. We found there is a high mobility among these eight categories, showing that labor markets in Mexico have been unstable in the last quarter century. A more precise analysis is done by dividing the population into three stages of life course: youth (15–24 years of age), early adulthood (25–44 years), and mature adulthood and old age (45–79 years). 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France, Italy, and Spain are among the leaders, and their experience can serve as inspiration for the establishment of social initiatives in Slovakia. They prefer social objective before making a profit; they are democratically organized and based on the unmet demand of the local community. The social enterprises are often creating sustainable jobs, bringing innovative solutions to social problems, and they have close links to active labor market policies. 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As set out in the decision of the Constitutional Court of RS, there is no doubt that the legislature has to adopt the law to regulate the attorneys? profession, but in its attempt of such regulation, the legislature always collides with the constitutional requirement that the attorneys are autonomous and independent. Therefore, when regulating the attorneys? profession, the legislature has to address the issues of autonomy and independence, and thus also the question of determining attorney tariffs, while certain issues must be left to be autonomously regulated within the attorneys? profession. The attorney tariff does not determine a mandatory price of legal services because an attorney and his client may always reach an agreement that is different from the attorney tariff. However, the tariff is binding when the court decides on the obligation to reimburse the party who has succeeded in a dispute for their attorney costs. 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He is the founder of The IEEE IWOBI conference series and the president of its Steering Committee, as well as the founder of both the InnoEducaTIC and APPIS conference series. He is an evaluator of project proposals for the European Union (H2020), Medical Research Council (MRC, UK), Spanish Government (ANECA, Spain), Research National Agency (ANR, France), DAAD (Germany), Argentinian Government, and the Colombian Institutions. He has been a reviewer in different indexed international journals (<70) and conferences (<250) since 2001. He has been a member of the IASTED Technical Committee on Image Processing from 2007 and a member of the IASTED Technical Committee on Artificial Intelligence and Expert Systems from 2011. \n\nHe has held the general chair position for the following: ACM-APPIS (2020, 2021), IEEE-IWOBI (2019, 2020 and 2020), A PPIS (2018, 2019), IEEE-IWOBI (2014, 2015, 2017, 2018), InnoEducaTIC (2014, 2017), IEEE-INES (2013), NoLISP (2011), JRBP (2012), and IEEE-ICCST (2005)\n\nHe is an associate editor of the Computational Intelligence and Neuroscience Journal (Hindawi – Q2 JCR-ISI). He was vice dean from 2004 to 2010 in the Higher Technical School of Telecommunication Engineers at ULPGC and the vice dean of Graduate and Postgraduate Studies from March 2013 to November 2017. 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He has (co)authored more than 150 publications in indexed journals, international conferences and book chapters, 1 book (in Greek), 3 edited books, and 5 journal special issues. His publications have more than 2100 citations with h-index 27 (GoogleScholar). His research interests include computer/machine vision, machine learning, pattern recognition, computational intelligence. \nDr. Papakostas served as a reviewer in numerous journals, as a program\ncommittee member in international conferences and he is a member of the IAENG, MIR Labs, EUCogIII, INSTICC and the Technical Chamber of Greece (TEE).",institutionString:null,institution:{name:"International Hellenic University",institutionURL:null,country:{name:"Greece"}}},editorTwo:null,editorThree:null},{id:"25",title:"Evolutionary Computation",coverUrl:"https://cdn.intechopen.com/series_topics/covers/25.jpg",isOpenForSubmission:!0,editor:{id:"136112",title:"Dr.",name:"Sebastian",middleName:null,surname:"Ventura Soto",slug:"sebastian-ventura-soto",fullName:"Sebastian Ventura Soto",profilePictureURL:"https://mts.intechopen.com/storage/users/136112/images/system/136112.png",biography:"Sebastian Ventura is a Spanish researcher, a full professor with the Department of Computer Science and Numerical Analysis, University of Córdoba. 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Main aspects of the topic are: Applying bioinformatics in drug discovery and development; Bioinformatics in clinical diagnostics (genetic variants that act as markers for a condition or a disease); Blockchain and Artificial Intelligence/Machine Learning in personalized medicine; Customize disease-prevention strategies in personalized medicine; Big data analysis in personalized medicine; Translating stratification algorithms into clinical practice of personalized medicine.",annualVolume:11403,isOpenForSubmission:!0,coverUrl:"https://cdn.intechopen.com/series_topics/covers/7.jpg",editor:{id:"351533",title:"Dr.",name:"Slawomir",middleName:null,surname:"Wilczynski",fullName:"Slawomir Wilczynski",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0033Y000035U1loQAC/Profile_Picture_1630074514792",institutionString:null,institution:{name:"Medical University of Silesia",institutionURL:null,country:{name:"Poland"}}},editorTwo:null,editorThree:null,editorialBoard:[{id:"5886",title:"Dr.",name:"Alexandros",middleName:"T.",surname:"Tzallas",fullName:"Alexandros Tzallas",profilePictureURL:"https://mts.intechopen.com/storage/users/5886/images/system/5886.png",institutionString:"University of Ioannina, Greece & Imperial College London",institution:{name:"University of Ioannina",institutionURL:null,country:{name:"Greece"}}},{id:"257388",title:"Distinguished Prof.",name:"Lulu",middleName:null,surname:"Wang",fullName:"Lulu Wang",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002bRX6kQAG/Profile_Picture_1630329584194",institutionString:null,institution:{name:"Shenzhen Technology University",institutionURL:null,country:{name:"China"}}},{id:"225387",title:"Prof.",name:"Reda",middleName:"R.",surname:"Gharieb",fullName:"Reda Gharieb",profilePictureURL:"https://mts.intechopen.com/storage/users/225387/images/system/225387.jpg",institutionString:"Assiut University",institution:{name:"Assiut University",institutionURL:null,country:{name:"Egypt"}}}]},{id:"8",title:"Bioinspired Technology and Biomechanics",keywords:"Bioinspired Systems, Biomechanics, Assistive Technology, Rehabilitation",scope:'Bioinspired technologies take advantage of understanding the actual biological system to provide solutions to problems in several areas. Recently, bioinspired systems have been successfully employing biomechanics to develop and improve assistive technology and rehabilitation devices. The research topic "Bioinspired Technology and Biomechanics" welcomes studies reporting recent advances in bioinspired technologies that contribute to individuals\' health, inclusion, and rehabilitation. Possible contributions can address (but are not limited to) the following research topics: Bioinspired design and control of exoskeletons, orthoses, and prostheses; Experimental evaluation of the effect of assistive devices (e.g., influence on gait, balance, and neuromuscular system); Bioinspired technologies for rehabilitation, including clinical studies reporting evaluations; Application of neuromuscular and biomechanical models to the development of bioinspired technology.',annualVolume:11404,isOpenForSubmission:!0,coverUrl:"https://cdn.intechopen.com/series_topics/covers/8.jpg",editor:{id:"144937",title:"Prof.",name:"Adriano",middleName:"De Oliveira",surname:"Andrade",fullName:"Adriano Andrade",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002bRC8QQAW/Profile_Picture_1625219101815",institutionString:null,institution:{name:"Federal University of Uberlândia",institutionURL:null,country:{name:"Brazil"}}},editorTwo:null,editorThree:null,editorialBoard:[{id:"49517",title:"Prof.",name:"Hitoshi",middleName:null,surname:"Tsunashima",fullName:"Hitoshi Tsunashima",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002aYTP4QAO/Profile_Picture_1625819726528",institutionString:null,institution:{name:"Nihon University",institutionURL:null,country:{name:"Japan"}}},{id:"425354",title:"Dr.",name:"Marcus",middleName:"Fraga",surname:"Vieira",fullName:"Marcus Vieira",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0033Y00003BJSgIQAX/Profile_Picture_1627904687309",institutionString:null,institution:{name:"Universidade Federal de Goiás",institutionURL:null,country:{name:"Brazil"}}},{id:"196746",title:"Dr.",name:"Ramana",middleName:null,surname:"Vinjamuri",fullName:"Ramana Vinjamuri",profilePictureURL:"https://mts.intechopen.com/storage/users/196746/images/system/196746.jpeg",institutionString:"University of Maryland, Baltimore County",institution:{name:"University of Maryland, Baltimore County",institutionURL:null,country:{name:"United States of America"}}}]},{id:"9",title:"Biotechnology - Biosensors, Biomaterials and Tissue Engineering",keywords:"Biotechnology, Biosensors, Biomaterials, Tissue Engineering",scope:"The Biotechnology - Biosensors, Biomaterials and Tissue Engineering topic within the Biomedical Engineering Series aims to rapidly publish contributions on all aspects of biotechnology, biosensors, biomaterial and tissue engineering. We encourage the submission of manuscripts that provide novel and mechanistic insights that report significant advances in the fields. Topics can include but are not limited to: Biotechnology such as biotechnological products and process engineering; Biotechnologically relevant enzymes and proteins; Bioenergy and biofuels; Applied genetics and molecular biotechnology; Genomics, transcriptomics, proteomics; Applied microbial and cell physiology; Environmental biotechnology; Methods and protocols. Moreover, topics in biosensor technology, like sensors that incorporate enzymes, antibodies, nucleic acids, whole cells, tissues and organelles, and other biological or biologically inspired components will be considered, and topics exploring transducers, including those based on electrochemical and optical piezoelectric, thermal, magnetic, and micromechanical elements. Chapters exploring biomaterial approaches such as polymer synthesis and characterization, drug and gene vector design, biocompatibility, immunology and toxicology, and self-assembly at the nanoscale, are welcome. Finally, the tissue engineering subcategory will support topics such as the fundamentals of stem cells and progenitor cells and their proliferation, differentiation, bioreactors for three-dimensional culture and studies of phenotypic changes, stem and progenitor cells, both short and long term, ex vivo and in vivo implantation both in preclinical models and also in clinical trials.",annualVolume:11405,isOpenForSubmission:!0,coverUrl:"https://cdn.intechopen.com/series_topics/covers/9.jpg",editor:{id:"126286",title:"Dr.",name:"Luis",middleName:"Jesús",surname:"Villarreal-Gómez",fullName:"Luis Villarreal-Gómez",profilePictureURL:"https://mts.intechopen.com/storage/users/126286/images/system/126286.jpg",institutionString:null,institution:{name:"Autonomous University of Baja California",institutionURL:null,country:{name:"Mexico"}}},editorTwo:null,editorThree:null,editorialBoard:[{id:"35539",title:"Dr.",name:"Cecilia",middleName:null,surname:"Cristea",fullName:"Cecilia Cristea",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002aYQ65QAG/Profile_Picture_1621007741527",institutionString:null,institution:{name:"Iuliu Hațieganu University of Medicine and Pharmacy",institutionURL:null,country:{name:"Romania"}}},{id:"40735",title:"Dr.",name:"Gil",middleName:"Alberto Batista",surname:"Gonçalves",fullName:"Gil Gonçalves",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002aYRLGQA4/Profile_Picture_1628492612759",institutionString:null,institution:{name:"University of Aveiro",institutionURL:null,country:{name:"Portugal"}}},{id:"211725",title:"Associate Prof.",name:"Johann F.",middleName:null,surname:"Osma",fullName:"Johann F. 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