Protein with anti-inflammatory properties produced in different strains of bacteria.
\r\n\t
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Venkateswarlu",coverURL:"https://cdn.intechopen.com/books/images_new/371.jpg",editedByType:"Edited by",editors:[{id:"58592",title:"Dr.",name:"Arun",surname:"Shanker",slug:"arun-shanker",fullName:"Arun Shanker"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"878",title:"Phytochemicals",subtitle:"A Global Perspective of Their Role in Nutrition and Health",isOpenForSubmission:!1,hash:"ec77671f63975ef2d16192897deb6835",slug:"phytochemicals-a-global-perspective-of-their-role-in-nutrition-and-health",bookSignature:"Venketeshwer Rao",coverURL:"https://cdn.intechopen.com/books/images_new/878.jpg",editedByType:"Edited by",editors:[{id:"82663",title:"Dr.",name:"Venketeshwer",surname:"Rao",slug:"venketeshwer-rao",fullName:"Venketeshwer Rao"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"4816",title:"Face Recognition",subtitle:null,isOpenForSubmission:!1,hash:"146063b5359146b7718ea86bad47c8eb",slug:"face_recognition",bookSignature:"Kresimir Delac and Mislav Grgic",coverURL:"https://cdn.intechopen.com/books/images_new/4816.jpg",editedByType:"Edited by",editors:[{id:"528",title:"Dr.",name:"Kresimir",surname:"Delac",slug:"kresimir-delac",fullName:"Kresimir Delac"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"59437",title:"Music and Brain Plasticity: How Sounds Trigger Neurogenerative Adaptations",doi:"10.5772/intechopen.74318",slug:"music-and-brain-plasticity-how-sounds-trigger-neurogenerative-adaptations",body:'\nGoing to concerts, listening to music in your living room, singing together or even playing an instrument is part of most people’s everyday life. Recent research indicates that apart from just changing the current mood this may have long-lasting influences on the brain. This contribution, therefore, describes how music shapes the brain as the outcome of interactions with the sounds. These interactions can be multifarious, as in the case of performing, listening or mentally imaging music, but they all show complex and widespread activity in many areas of the brain. This activity, moreover, is related to training, previous exposure, personal preference, emotional involvement and many other modulating factors related to the cultural background and biological repertoire of each individual [1, 2, 3, 4, 5, 6, 7]. Musical training, moreover, is related to structural changes within auditory and motor areas of the brain and reinforces functional coupling of these regions during musical tasks as evidenced by many neuroimaging studies [8, 9, 10]. These changes have been observed also in white-matter tracts, such as the corpus callosum, the corticospinal tract and the arcuate fasciculus [11, 12, 13]. Studies (particularly those with a longitudinal design) showing the causal relation between the brain changes and the duration of musical training have convinced some researchers to consider musical training as a model for investigating practice-related brain plasticity in humans [14].
\nMusic is a powerful stimulator of the brain. Acoustically, it consists of time-varying sound events that are characterised by a large number of features—more than hundred features can be computationally extracted that are tracked by several regions of the brain [15]. Many low-level features, such as timbre and pitch, are partly processed in Heschl’s gyrus and the right anterior part of the superior temporal gyrus, in which the primary and non-primary auditory cortices are located [16, 17]. Besides auditory cortices, also motor regions, such as the supplementary motor area and the cerebellum, are involved during musical activities, including both playing and listening. Due to audio-motor coupling that is necessary for playing an instrument, listening is influenced by the motor demands intrinsic to musical practice, even to the extent that this would become manifest also in the brain responses to music listening alone [18, 19]. Moreover, practising and performing music is a complex, multimodal behaviour that requires extensive motor and cognitive abilities. It relies on immediate and accurate associations between motor sequences and auditory events leading to multimodal predictions [10, 20, 21], which engage broad networks of the brain [16, 22, 23]. Music training has thus been associated with changes in the brain, and some of these changes have been causally linked to the duration of the training, which makes the musician’s brain a most interesting model for the study of neuroplasticity [9, 24]. This holds in particular for performing musicians, who provide a unique pool of subjects for investigating both the features of the expert brain and, when considering the length of the training, also the neural correlates of skill acquisition. Musicians’ training and practice require the simultaneous integration of multimodal sensory and motor information in sensory and cognitive domains, combining skills in auditory perception, kinaesthetic control, visual perception and pattern recognition [25, 26]. In addition, musicians have the ability to memorise long and complex bimanual finger sequences and to translate musical symbols into motor sequences (see Figure 1). Some musicians are even able to perceive and identify tones in the absence of a reference tone, a rare ability termed absolute pitch [27, 28].
\nIllustration of several perceptual, motor, interoceptive and emotional skills that are acquired during musical training.
The brain changes that musical training entails are numerous and well-documented [2, 3, 5, 9, 24, 26, 27, 28, 29, 30, 31, 32, 33, 34]: they involve brain regions important for auditory processing, coordination of fast movements and cognitive control, as well as sensory-to-motor coupling mechanisms (see [35], for an overview). While some of these changes might be what characterise individuals that decide to undertake a musical profession, and hence might exist at birth, others could be a direct result of training, as suggested by the significant relations between years of training and brain measures (e.g., [36]), as well as by longitudinal designs recording brain responses before and after music training (e.g., [29, 37]).
\nHere, we propose that the differences observed between the brains of musicians and non-musicians can be attributed to neuroplastic adaptations responding to the challenging demands of musical practice. Alternative explanations are also possible, such as that the differences exist even before training in those individuals that choose music as their profession, but accumulating evidence points at a causal relation between music training and brain changes. The behavioural correlates of these differences are multiple and can be seen especially in childhood (e.g., [38]). Besides, it has been shown that music may be beneficial in relation to a number of symptoms in several kinds of impairment, such as epilepsy, Alzheimer’s disease, Parkinson’s disease and senile dementia (see [39] for an overview). Hence, it is possible to conceive of dealing with music in educational, clinical and therapeutic terms.
\nIn this contribution, we first expose the concept of adaptation, both from the phylogenetic and ontogenetic points of view. We then narrow down this concept by putting forward the hypothesis of music-induced neuroplasticity, with a first distinction between macrostructural and microstructural adaptations. Thereafter, we consider the reorganisation of the brain as the outcome of learning and skill acquisition, both at a structural and functional level of description with a major focus on the adult musician or listener as a model for the interaction between ontogeny and phylogeny. This latter, further, is considered from the point of view of network science with a major focus on the role of resting-state networks. Clinical and therapeutic applications, finally, are envisioned also.
\nBrain plasticity is an adaptation to the environment with an evolutionary advantage. It allows an organism to be changed in order to survive in its environment by providing better tools for coping with the world [40]. This biological concept of adaptation can be approached from two different scales of description: the larger evolutionary scale of the human as a species (phylogeny) and the more limited scale of the human from newborn to old age (ontogeny). This phylogenetic/ontogenetic distinction is related to the “nature/nurture” and “culture/biology” dichotomy, which refers to the neurobiological claims of wired-in circuitry for perceptual information pickup as against the learned mechanisms for information processing and sense-making and immersion in a culture [41, 42].
\nThese approaches may seem to be diverging at first glance, but they are complementary to some extent. This holds, in particular, for the here-hypothesised music-induced plasticity, which espouses a biocultural view that aims at a balance between genetic or biological constraints and historical/cultural contingencies. This places all human beings on equal ground (unity) by stating that diversity in culture is only an epiphenomenon of an underlying biological disposition that is shared by people all over the world [43]. The assumed unity is attributed to the neural constraints that underlie musical processing in general, but these constraints should not be considered as a static dispositional machinery. The picture that emerges from recent research is arguing, on the contrary, for a definition of the neural machinery as a dynamic system that is able to adapt in answer to the solicitations of a challenging environment [6]. The neurobiological approach to music, therefore, deals not only with the nature and evolution of the innate and wired-in neural mechanisms that are the hallmark of the hominid phylogenetic evolution but also with the ontogenetic development of these mechanisms [43]. As such, it makes sense to conflate neurobiological and developmental claims by taking the concept of adaptation as a working hypothesis.
\nThe relation between adaptation and development, however, is asymmetrical in the sense that it is possible to conceive of development without adaptation, but no adaptation is conceivable without development. This development, further, can be natural, when it is the outcome of maturation, but it is possible also to intervene in its trajectory by combining development and learning. This is the case when an organism faces continued and long-term exposure to challenging environments, which triggers plastic changes in the structure and the functions of the brain. This brings us to the concept of brain plasticity, which refers to the fact that neuronal circuits are tuned in close interaction with the environment. It was introduced by William James, who defined plasticity as “the possession of a structure weak enough to yield to an influence, but strong enough not to yield all at once” [44] (p. 106). The idea was further developed by Ramón y Cajal, who claimed that to fully understand the phenomenon it is necessary to admit the formation of new pathways in the brain through ramification and progressive growth of the dendritic arborisation and the nervous terminals in addition to the reinforcement of pre-established organic pathways. The same idea was elaborated further by Donald Hebb, who proposed that neuronal cortical connections are strengthened and remodelled by experience. There is, however, another aspect of plasticity that goes beyond the level of synapses and that incorporates the level of cortical representation areas or cortical maps, which can be modified by sensory input and training [45]. It is suggested, in this regard, that additional neurons are recruited when they are needed and that rapid and transient alterations of cortical representations can be seen during learning tasks. Such short-term modulations are important in the acquisition of new skills, but they can lead also to structural changes in the intra-cortical and sub-cortical network once the skill has been established.
\nThe evolutionary claims of adaptation—both at the phylogenetic and ontogenetic level—have received empirical evidence from neuroimaging and morphometric studies. In order to elucidate its underlying mechanisms, there is currently a whole body of research related to the psychobiological approach to the study of action, cognition and perception. A major claim in this research field is that the nervous system provides the immediate, necessary and sufficient mechanisms that underlie all mental processes, and that mental processes are reducible to the function, arrangement and interaction of neurons as the constituent building blocks of the nervous system [46]. This is the axiom of psychobiological equivalence, which claims an equivalence of maintained information from the neural to the psychological state [47]. The related research revolves around three major themes: (i) the localisation of functions in the brain, (ii) the representation or coding and (iii) the dynamic change or learning [46]. The first investigates which brain structures are responsive for particular processes. The second investigates how neural networks represent, encode or instantiate cognitive processes, both at macrostructural and microstructural level of description (see [48]). The third, finally, investigates how our brain adapts to experience and learning, what changes occur in its neural networks and how these changes correspond to externally observed behaviour.
\nThe third theme—dynamic change or learning—concerns the neural correlates of skill acquisition and has been studied mainly at the level of perceptual processing and motor output. Yet, there is also the whole domain of creativity [49], musical aesthetics [6, 50, 51, 52, 53] and human interaction [54, 55], which have been poorly investigated in relation to long-term music training. However, the topic is exemplary of a paradigm shift in current neuromusicological research, with a transition from a static conception of brain modules to a conception of reorganisational plasticity of the developing and adult brain [6]. Plasticity, in fact, is a fundamental organisational feature of the human brain, which can be modified throughout the life span in response to changes in environmental stimulation. This has been observed not only during a critical period in the developing brain but also even throughout the whole life span.
\nSkill learning, such as learning to play a musical instrument, can thus be used for the study of neuroplasticity. It typically starts early in life, while the brain is most sensitive to plastic changes, and continues often throughout life. It involves multiple sensory modalities and motor planning, preparation and execution systems [27, 56]. The role of environmental enrichment—being defined as a combination of complex inanimate and social stimulation [57]—on the other hand, should be stressed also as an emerging area of research. Music, in this view, can be considered as “sounding environment” [42], which is likely to drive brain plasticity. Even in the foetal phase of development, sounds can trigger ways of implicit learning [58]. Neonates and infants also learn to talk and sing quasi-effortless as the result of mere exposure, thus demonstrating implicit learning and developmental plasticity, which is even cross-modal to a large extent, in the sense that loss of one sensory modality may lead to neural organisation of the remaining modalities [2].
\nNeuroplasticity is also related to the field of sensory-motor learning, with a major role attributed to the challenges of a rich environment. It favours multiple interactions with the world—both at the sensory and at the motor level—stressing the interdependency of an organism and its environment through which it “enriches its repertory of genetic adaptations with acquired dispositions that are immediately at hand and mobilizable when confronted with a situation that can be foreseen or recognized as a familiar one” [59] (p. 925).
\nMusical training, accordingly, may be related to sensory and motor changes in the human brain of professional musicians. As a rule, music training involves years of sensory-motor training, often beginning in early childhood, with the aim to develop an expertise in a chosen instrument or mastery over the own voice, together with an improvement of the ability to attend to the fine-grained acoustics of musical sounds, including pitch, timing and timbre [3]. The brains of musicians might adapt to the demands of their instrumental practice at two levels: the gross anatomical differences between professional musicians and amateurs or laymen, and the subtle functional differences after enhanced musical practice and/or experience, which have to be sought in ever finer modifications of synaptic strength in distributed cortical networks. As such, it is possible to distinguish between macrostructural and microstructural adaptations. The macrostructural differences related to volume, morphology, density and connectivity of brain structures are measured with magnetic resonance imaging (MRI), whereas the microstructural differences in the functional activity of brain regions are measured with functional magnetic resonance imaging (fMRI), positron emission tomography (PET) and neurophysiology (electroencephalography, EEG and magnetoencephalography [MEG]) (see [5] for an overview). It is further hypothesised that functional reorganisation may cause structural adaptation. For instance, bimanual instrument training, such as for the piano, may cause an increase in cortical functionality for symmetric areas involved in motor, auditory and visuo-spatial processing, as well as in the white matter tracts of the corpus callosum [20] as compared with less bimanual training (such that for the violin) and especially as opposed to laypersons.
\nAt the macrostructural level, studies show differences in the size of the primary motor cortex size, the cerebellum, the planum temporale, the corpus callosum, Heschl’s gyrus and the arcuate fasciculus, all of which seem to correlate with the ability of musicians to identify and process acoustic variations [60]. The microstructural adaptations, on the other hand, happen at the level of individual neurons and synapses, with the aim to change the efficacy of neural connectivity. In general, the brain shows adaptation to extraordinary challenges by giving birth to new neurons (neurogenesis) and glial cells and by the formation and remodelling of new connections by the outgrowth of dendrites, axonal sprouting and increasing or strengthening of synaptic connections [61] (see Box 1 for an overview). Such adaptations have been studied in the context of deafferentation studies—in the case of brain lesion—and in the case of motor skill learning [62]. In the former, some cortical remodelling has been found, including microstructural changes, such as the strengthening of existing synapses, the formation of new synapses (synaptogenesis), axonal sprouting and dendrite growth. In the latter, similar changes have been found, including an increased number of synapses per neuron and changes in the number of microglia and capillaries [63], which all lead to volumetric changes that are detectable also at the macrostructural level [27, 64].
\nOverview of some basic mechanisms for refinement of the neural circuitry.
\nMyelinisation: the acquisition, development or formation of a myelin sheath around a nerve fibre. This fatty coating serves as insulation of individual fibres to enhance specificity of connections and increases markedly the quick and accurate transmission of electrical current from one nerve cell to another.
\nPruning: a process that helps sculpt the adult brain and by which neurons and synaptic connections that are no longer used or useful are eliminated in order to increase the efficiency of neuronal transmissions.
\nSprouting: a process by which a neuron generates additional branches or outgrowths to establish new links between existing neurons, as seen frequently in the case of growth of axons or dendrites from a damaged or intact neuron that projects to an area that is denervated.
\nSynaptic plasticity: strengthening or weakening of the synaptic links either by modulating the strength of synapses or by altering their numbers.
\nSynaptic efficacy: changes in the number or properties of postsynaptic receptors in transmitter release and the formation of new synapses (synaptogenesis).
\nAdult neurogenesis: birth of neurons from neural stem cells in the adult brain. In humans, adult neurogenesis has been shown to occur only in the hippocampus (particularly in the sub-granular zone of the dentate gyrus) and in the striatum. It differs from developmental neurogenesis.
\nThe bulk of studies on neuroplasticity has been performed in the context of within-modality plasticity, particularly in the domain of sensory and motor modalities. They aim at demonstrating the adaptive capabilities of the human brain to shape the processing of sensory stimuli or to perform motor acts after repeated sensory exposure or action [9]. With regard to the sensory modality, animal studies have shown that environmental change critically affects brain development. Experience-driven neural activity, in fact, regulates the refinement of the neural circuitry by influencing various neural processes, such as synapse formation, pruning and synaptic plasticity (see Box 1) with modifications in synaptic connectivity as a result [65]. This enhanced connectivity, further, acts as a basis for learning and memory through alterations at the level of neural circuits [66], such as strengthening or weakening of the synaptic links or altering their number, changes in the number or properties of postsynaptic receptors in transmitter release and the formation of new synapses. The result is an increase in synaptic strength, which may be persistent and facilitate learning and memory so that experience-dependent plasticity could involve selective changes in pre-existing brain circuits [65].
\nAs to the enhanced auditory skills, it has been argued that they may prime the brain for the processing of musical sounds and that these skills may percolate to other domains, such as speech, emotion and auditory processing in general [67, 68]. This has been observed already in the early stages of the auditory pathway, which are located mainly in the brainstem. Musicians have enhanced temporal and frequency coding in the auditory brainstem with earlier (as early as 10 ms after acoustic onset) and larger responses than non-musicians to both speech and music stimuli. This has been shown for the onset response and the frequency-following response (FFR), i.e., a neuronal ensemble response that phase-locks to the incoming stimulus and that underlies perception of pitch as it relates to the sustained portion of a periodic sound with less or more stable frequencies [68, 69].
\nThe role of auditory brainstem processing of behaviourally relevant sounds such as speech and music is important here. It can be measured by using the onset response and the FFR to see how the brainstem represents pitch, timing and timbre [68]. It has been shown that both temporal and spectral characteristics of sounds are preserved in this subcortical response (see [70] for an overview), reflecting the physical properties of sound with an unrivalled fidelity. As a rule, it occurs automatically at pre-attentive levels of auditory processing but is shaped by both long-term and short-term experience [71, 72, 73]. Subcortical function, moreover, is neither passive nor hardwired but interacts dynamically with higher-level cognitive processes refining the transcription of sounds into neural code. Hence, the responses do not originate merely in the brainstem but receive feedback from top-down cortical influences even at the earliest stages of auditory processing [3] via corticofugal feedback pathways [74, 75]. As such, it can be demonstrated that musical practice changes the early sensory encoding of auditory stimuli [68] relying on a top-down feedback system—consisting of efferent effects on cochlear biomechanics—that is continuously and automatically engaged to extract and represent regularities in the auditory system [3]. Musical training is thus not limited to the modification of cortical organisation but the modifications extend to subcortical sensory structures and generalise to early processing of speech and sounds in general.
\nMoreover, early auditory evoked responses and particularly the negative–positive complex (N19-P30) in the auditory evoked potential [76] localised in the primary auditory cortex (the anteromedial portion of Heschl’s gyrus) have been found to be larger in musicians compared to amateurs and non-musicians. Moreover, it has been found that the generating neural tissue, namely the grey matter volume of the primary auditory cortex, was broader in volume for professional musicians [77] as compared to laypersons. It thus seems that music can trigger both macrostructural and microstructural or functional changes, not as separate and distinct levels of adaptations, but as phenomena that are dynamically and tightly interconnected.
\nMusic can trigger plastic changes in the brain, as evidenced by the rich history of structural and functional neuroimaging studies of the past decades. Recent advances in functional neuroimaging have furthermore provided new tools for measuring the functional interactions and communication between distinct regions in the brain and for examining their functional connectivity [78]. In an attempt to study the brain as a complex network of functionally and structurally interconnected regions, a fuller understanding of its organisation and function is proposed by relying on the contributions of network science [79], which investigates complex systems in terms of their elements and the relationships and interactions between these elements.
\nFunctional connectivity can be defined as the temporal dependence of neuronal activity patterns of anatomically separated and removed regions in the brain, reflecting the level of communication between them [80]. It makes it possible to examine the brain as an integrative network of functionally interacting regions and to gain new insights into large-scale neuronal communication in the human brain. Such whole-brain connectivity patterns can be studied by measuring the synchronisation of spontaneous fMRI or MEEG time-series reflecting neural activity of anatomically separated brain regions, which are recorded during rest. These resting–state networks are believed to reflect the functional communication between brain regions [78, 81] and suggest an ongoing information processing and functional connectivity between them even at rest, which is related to neuronal firing. The pattern of correlations between distinct brain areas, moreover, points at the existence of organisational networks in the brain [81], which seems to be analogous to the networks that are engaged during the performance of sensory-motor and cognitive tasks, and which are dependent upon the brain’s anatomical connectivity [10]. Such spontaneous neuronal interaction has been first investigated in motor cortices but were later extended to other cortical systems, such as the visual and auditory networks, the default mode network (DMN) and attention and memory related regions. It has been suggested that at least 10–12 resting-state networks (RSNs) can be detected in the cerebral cortex in resting state, which implicates that they represent some intrinsic form of brain connectivity with temporal correlations between spatially discrete regions [82].
\nDMN has been related to specific brain functions, such as self-referential thoughts, emotional perspectives and levels of self-awareness. DMN is believed to be a neural circuit that constantly monitors the sensory environment and displays high activity during lack of focused attention on external events [83]. It seems to function as a toggle switch between outwardly focused mind states and the internal or subjective sense of self [84] and can be used to explore the functional connections of the complex integrative network of functionally linked brain regions, which continuously share information with each other. As such, there are interconnected resting-state neuronal communities or functional brain networks with functional communication between them. Being organised according to an efficient topology, they combine efficient local information processing with efficient global information integration with the most pronounced functional connections found between those regions that share common functions.
\nOverall, resting-state fMRI oscillations reflect ongoing functional communication between distinct brain regions [78], which makes them indicative of the level of cognitive functioning in general. There seems to be, in fact, a link between an efficient organisation of the brain network and intellectual performance—this is the neural efficiency hypothesis—so that functional connectivity patterns may be used as a powerful predictor for cognitive performance [85]. This resting-state connectivity, further, is not to be considered as an established and fixed property, but as a state that can be modulated by recent experiences and learning episodes, both within and between the networks they recruit. Such modulation points in the direction of a learning consolidation function of resting-state brain activity, as evidenced by the findings that high learners manifest stronger pre-task resting-state functional connectivity between the involved regions than low learners [10]. It thus seems that, even in the absence of external stimuli or demands, the brain is constantly sharing information. It thus consolidates recent learning and maintains the association of activity of brain areas that are likely to be used together in future [86].
\nInitial research suggests that musical training might enhance this pattern of increased resting-state connectivity by triggering heightened connections at a functional level between those brain regions that are structurally and functionally altered as the result of training. This is manifested even during a task-free condition, pointing to the “silent” imprint of musical training on the human brain [35]. Research on the differences between musicians and non-musicians in their functional connectivity during rest, however, is still in its infancy [10, 82]. By selecting predefined seed regions for computing connectivity analysis, increased connectivity between contralateral homologue regions has been found in musicians between prefrontal, temporal, inferior-parietal and premotor areas [35]. It is to be questioned, however, whether the study of predefined regions or seed regions does not neglect residual whole-brain dynamics. However, for the seed regions for which plastic changes in musicians have been found already—as evidenced by increased grey matter volume—connectivity analyses have revealed brain areas whose resting-state time series activity was more closely synchronised with one of them. Four networks were found to supply integrative interpretations for the cognitive functions during musical practice: (i) autobiographical memory-related regions belonging to the default mode network, recruited by the encoding, storage and recall of melodies with an emotional and biographical quality; (ii) areas that belong to the salience network with access to semantic memory that is related to the storage of music in terms of verbal labels and auditory structure; (iii) regions that are implied in language processing and the resting-state auditory network and (iv) structures that belong to the executive control network, and which could subserve the motor modulation required for an emotionally expressive interpretation of music. The question whether this practice-related plasticity is triggered by local grey matter volume, however, is not yet satisfactorily resolved, in the sense that other variables may be implicated in the expertise-related resting-state functional reorganisation of musician’s plastic brain [10].
\nTo stretch further our hypothesis about music-induced neuroplastic adaptation, music, as a cognitive-demanding activity stimulating neuroplasticity, may be able to slow down, arrest or even reverse the detrimental effects of ageing on learning and memory capacity of the elderly [33]. Recent studies have provided evidence that music-induced plasticity may help also to overcome neurological impairments, such as neurodevelopmental disorders and acquired brain injuries [56]. For instance, attentive music listening recruits multiple forms of working memory, attention, semantic processing, target detection and motor function, relying mainly on bilateral brain areas—superior temporal gyrus, intraparietal sulcus, precentral sulcus, inferior sulcus and gyrus, and frontal operculum—which all serve general functions rather than music-specific cortical regions [87, 88]. Complex musical tasks, moreover, engage the co-activation of many processes involving widely distributed and partly interchangeable substrates of the brain [89]. This may explain, to some extent, the sparing of some musical functions in cases of progressive destruction of some areas in degenerative diseases of the brain. This has been shown most typically in the case of Alzheimer’s disease (AD), which is characterised by a general and progressive decline in cognitive function, with the first symptom as an impaired episodic memory. Music, in this case, has been reported as one of the domains in which general skill and memory are preserved in spite of otherwise severe impairment [90]. This preserved musical processing, moreover, is not limited to procedural memory but often includes also stories of music, which can be used as an effective mnemonic device [91].
\nHence, music may shape the development of normal and healthy human beings over the lifespan, but its potential as a non-pharmacological interventional aid for caregivers to help the cognitive and emotional capacity of patients with neurological and psychiatric brain disorders is receiving growing interest [15]. The use of resting-state fMRI techniques, e.g., with a main focus on the default mode network, seems to be well-suited to examine possible functional disconnectivity effects in disorders such as Alzheimer’s disease, depression, dementia and schizophrenia. Also, other neurogenerative diseases like multiple sclerosis and amyotrophic lateral sclerosis seem to show changed connectivity in the default network as well as in other resting-state networks [78]. This may suggest that neurodegenerative diseases would attack interconnected cortical networks rather than single regions in the brain [92] and can thus be targets of a music intervention aimed at stabilising abnormal patterns of functional connectivity between compromised brain areas.
\nMusic has been used already as a treatment for some psychiatric and neurological pathologies, such as schizophrenic disorders, Alzheimer’s disease, Parkinson’s disease, cerebral ischemia, pain, autism, anxiety and depression [15]. Music, furthermore, has been reported to improve also the well-being and cognitive functions in healthy adults, such as autobiographical memory, semantic memory, language ability and cognitive functions, and to alleviate neuropsychiatric symptoms, such as agitation, apathy, depression and anxiety (see [39] for an overview). Effects of music on AD are exemplary of the mechanisms that might mediate the impact of music on human well-being. Latent benefits of musical mnemonics as an aid to standard mnemonic methods, which may seem to be insufficient for AD patients, have been reported (for a review, see [15]). The mechanisms behind these memory-enhancing effects, however, are still not fully understood, but there is strong evidence for a benefit of music as a mnemonic device in a variety of clinical settings [91]. A possible explanation is that the areas of the brain associated with music cognition are preferentially spared in the case of AD. It has been suggested that procedural memory and priming effects for musical stimuli remain intact, whereas short-term and long-term episodic memory for melodic excerpts is impaired [93].
\nThis dissociation between memory and general performance in AD patients holds in particular for listening to their favourite songs, which seems to recruit previously encoded memories. These memories seem to support and sustain brain introspection via connectivity within the default mode network and also to effectively reprocess autobiographic and episodic memories [84]. An additional explanation for this dissociation is that in patients with general cortical and hippocampal atrophy, which impairs standard episodic learning, musically-associated stimuli allow for a more diversified encoding. Music processing, in that case, encompasses a neural network that is recruiting from multiple areas of the brain, including cortical as well as subcortical areas. Musical stimuli and stimuli accompanied by music seem to create a more robust association at the stage of encoding and support a more composite encoding and retrieval process by inducing oscillatory synchrony in those neural networks that are associated with learning and memory [91, 94].
\nNeuroplasticity is now an established topic in music and brain studies. Revolving around the concept of adaptation, it has been found that the brain is able to adapt its structure and function to cope with the solicitations of a challenging environment. This concept can be studied in the context of music performance studies and long-term and continued musical practice. It has been shown that some short-term plastic changes can even occur in the case of merely listening to music—without actually performing—(e.g., [95]) and in the short-time perspective of both listening and performing (e.g., [96]). Attentive listening to music in a real-time situation, in fact, is very demanding: it recruits multiple forms of memory, attention, semantic processing, target detection and motor function [18, 97]. As such, we propose here that music represents a sort of enriched environment that invites the brain to raise its general level of conscious functioning.
\nTraditional research on musical listening and training, however, has focussed mainly on structural changes, both at the level of macro- and microstructural adaptations. This has been well-documented with morphometric studies, which aimed at showing volumetric changes of target areas in the brain as the outcome of intensive musical practice. Recent contributions, however, have shown that the brain can be studied also from the viewpoint of network science. The brain, in this view, is not to be considered as an aggregate of isolated regions, but as a dynamic system that is characterised by multiple functional interactions and communication between distinct regions of the brain. Whole-brain connectivity patterns can be studied by measuring the co-activation of separate regions. Much is to be expected from the study of resting-state networks with a special focus on the default mode network. These networks seem to be indicative of the level of cognitive functioning in general and are subject to the possibility of modulation by experience and learning, both in the developing and in the mature brain. We propose that music has the potential to alter the organisation of these brain networks and enhance the connectivity of the brain, both in normal people and in those with an impaired brain.
\nA major emerging topic, therefore, is the tension between neurogenerative and neurodegenerative forces with the critical question as to the possible role of music as an intervening force to develop, maintain or even restore the connectivity in brain tissue. The idea that age-related cognitive decline may be slowed, arrested or even reversed through appropriately designed training or activities, such as musical practice, is supported already by some research. Moreover, the finding that the adult brain can undergo continual modifications highlights the potential of music intervention for inducing the plastic changes that can ultimately attenuate the impairments due to brain injury. Much more research, however, is still needed towards an integration of findings from neuroscience, education, music therapy and development.
\nThe human gastrointestinal tract is formed by a complex ecosystem which includes the gastrointestinal epithelium, immune cells, and resident microbiota [1] and comprehends one of the biggest existent interfaces between the host, environmental factors, and antigens in the human body.
\nThe intestine encompasses a broad variety of microorganisms (bacteria, archaea, eukarya, and viruses) [2] from more than 3500 different species [3, 4] that coevolved with the host in a mutually beneficial relationship [5, 6]. The composition and density of bacterial populations in adult individuals differ considerably over the GIT. The area of the GIT that has highest microorganism abundance is the colon (1014) followed by dental plaque (1012), ileum (1011), saliva (1011), and skin (1011) [7]. However, low concentrations (up to 102–107 cells/mL) and bacterial diversity are found in the upper GIT (stomach, duodenum, jejunum) [3, 4], since the presence of acid, bile salts, and pancreatic secretions hinders the bacterial colonization [8], so that there is no nutritional competition between the microbiota and the host [9]. Thus, both function and structure of microbial communities are significant and are closely related. However, function could be the more important measure of microbiome health, since bacterial ecology suggests that analogous ecosystems have similar function although they have moderately diverse composition [10, 11].
\nThe importance and the specific functions that gut microbiota has in human nutrition and health are well settled. The attributed functions can be classified in three classes: metabolic, protective, and trophic [12]. The gene diversity of the microbial community provides a variety of enzymes and biochemical pathways, specific to the host, able to contribute to short-chain fatty acid (SCFA) production by carbohydrate fermentation and production of some vitamins such as K, B12, biotin, folic acid, and pantothenate. These factors added to synthesis of amino acids from ammonia or urea contributing to the metabolic function of the microbiota [13, 14].
\nThe gut microbiota’s protective function is related to barrier effect, once the resident bacteria generate a resistance line which avoid pathogens/opportunistic bacteria and maintain normal mucosal function. The activity of some bacteria to secrete antimicrobial substances, such as bacteriocins, is able to inhibit the growth of other bacteria and nutrient competition [15, 16].
\nRegarding trophic functions of gut microbiota, the interaction between resident microorganisms has influence in differentiation and proliferation of epithelial cells [17], as well as in the development and regulation of the immune system by numerous and varied interactions between microbes, epithelium, and gut lymphoid tissues [18].
\nIt is important to highlight that the interactions between the gut microbiota and the host immune system are required to preserve the gut homeostasis [19, 20, 21]. When this relationship is affected, alterations in bacterial function and diversity lead to the imbalance in the composition of the resident microbiota, favoring either the growing of pathogenic bacteria or the decreasing in beneficial bacteria in a process known as dysbiosis [22], which appoint a great threat to gut integrity and is intrinsically related to the development and progression of several diseases, such as inflammatory bowel diseases.
\nOne of the most well-characterized chronic inflammatory diseases that mainly affect the digestive tract is inflammatory bowel disease (IBD), which includes ulcerative colitis (UC) and Crohn’s disease (CD). The exact etiology of IBD is still unclear, but the strict relation between genetic and the environmental factors, such as enteric immune dysregulation and alterations in the intestinal microbiome [23, 24], is broadly known. Besides, these diseases generate substantial morbidity and have a high prevalence in developed countries (5 in 1000 individual are affected) they remain to increase in developing nations [25].
\nBoth diseases, UC and CD, present different pathogenesis, symptomatology, inflammatory profiles, and gut microbiota composition. CD is characterized by the irregular transmural inflammation (extending deeply into the submucosal regions) which can affect any portion of the GIT and often made difficult by strictures, abscesses, and fistulae. On the other hand, the inflammation presented in UC is restricted to the superficial layers of the intestinal mucosa characterized by mucosa erosion and/or ulcer, generally localized in the region of the gut most colonized by bacteria, the colon [26, 27]. In addition, regarding the immune response associated with these diseases, it is possible to relate CD with an increased IL-12, IL-23, IL-27, interferon γ (IFN-γ), and tumor necrosis factor-α (TNF-α) production, all associated with Th1 and Th17 immune responses, different from UC which is correlated with a Th2 immune response, with high levels of IL-5 and transforming growth factor-β (TGF-β) production [28].
\nIt is important to highlight that the principal cause of death in IBD patients is colorectal cancer (CRC) [29]. Frequent episodes of inflammatory process in the intestinal mucosa are related to development of this disease, which is the second most frequently identified cancer in females and the third in males.
\nThere are increased evidences that environmental factors such as lifestyle and diet alterations have effect in CRC incidence [30]. This effect has been documented because there is evidence showing an essential relationship between dietary antigens and antigens of commensal bacteria with the regulatory T cells (Tregs), which maintain the immune tolerance and, consequently, reduce the risk of tumorigenesis associated with inflammation [31].
\nIn this context, it was reported that the higher consumption of diet rich in grains and vegetables decreases the incidence of CRC. This effect involves different mechanisms such as the diminution in the fecal transit time due to the increase in the stool bulk, and consequently, it reduces the contact of carcinogen with colon cells and the fermentation of these fibers of colonic components [14, 32]. In addition, significant reduction in concentration of acetate, propionate, and butyrate with increase in fecal pH [33] and the decrease in the number of obligate anaerobe microorganisms have been reported in individuals with colon cancer [34] when compared with healthy people. Thus, intestinal environmental alterations are the keys to evolution toward adenoma and afterward to CRC progression [35].
\nIt has been also reported that up to 30% of patients with UC need surgical management such as the restorative proctocolectomy with ileal pouch-anal anastomosis (IPAA) [36]. This procedure removes the entire colon and rectum while preserving the anal sphincter and, hence, normal bowel function and fecal continence, therefore acting as an internal pelvic place for intestinal contents [37]. Around 50–60% of UC patients with following IPAA develop inflammation in the ileal pouch, generating the condition called “pouchitis.” The reported incidence of pouchitis is variable, generally because of the diagnostic criteria that have been used to define this syndrome [38, 39]. In addition, although its pathogenesis is uncertain, the main hypothesis for the mechanism by which the disease occurs is the break in the mucosal barrier generated by dysbiotic microbiome in susceptible patients, generating an unusual mucosal immune activation [40]; still the disease typically responds to antibiotics.
\nCorresponding to the increased attention given to the role of the intestinal microbiota in a variety of diseases, there has been an intense exploration of potential means to manipulate the intestinal microbiome either by probiotic administration or fecal microbiota transplant (FMT) for therapeutic effect [41].
\nIn this context, a randomized clinical trial based on a 1-week treatment with anaerobically prepared donor FMT, compared with autologous FMT, resulted in a higher probability of remission in 8 weeks for patients with UC, revealing that stool administration from healthy donors to UC or CD patients is an intervention that seeks to restore a healthier balance of gut microbes and control IBD [42]. Data on FMT for Crohn’s disease is rather more limited than for UC, but it has been shown that single standardized FMT resulted in a clinical remission sustained for more than 9 months in CD patients [43]. However, the authors suggest that further studies are needed to enhance the knowledge about the use of stool transplantation for IBD treatment.
\nAlteration in the gut microbiome composition with increase in some groups of microorganisms, such as Clostridium and Fusobacterium, was also reported in patients with pouchitis [44, 45]. In this context, literature evidences indicate that the probiotic administration such as VSL#3 is effective in the chronic pouchitis prevention [46]. On the other hand, FMT to pouchitis treatment did not report the same beneficial results. Only three reports with this approach [47, 48, 49] exposed that neither clinical remission nor any adequate response was observed in the evaluated patients suggesting that the efficacy of FMT for pouchitis after proctocolectomy is limited [49]. The importance of standardization of this procedure needs to be highlighted to improve its efficacy, since frequency, route of administration (e.g., endoscopy, nasogastric tube, colonoscopy), and the criteria of choice of healthy donor are very important parameters to be considered.
\nDifferent chemotherapy regimens such as FOLFOX (5-fluorouracil and oxaliplatin), FOLFIRI (5-fluorouracil and irinotecan), and triple FOLFOXIRI regimen (5-fluorouracil, oxaliplatin, and irinotecan) [50, 51] are adopted for different types of cancer but with a broad range of collateral effects.
\nMucositis is the most common side effect in patients undergoing chemotherapy/radiotherapy treatments, which consist in an inflammation and/or ulcers in the gastrointestinal tract [52] with consequent loss of cells from the epithelial barrier of the GIT. Many symptoms are related to gastrointestinal mucositis, such as diarrhea, severe abdominal pain, bleeding, fatigue, malnutrition, dehydration, electrolyte imbalance, and infections, with potential fatal complications which can conduce to reduction or interruption of antitumor treatment [53] and consequently leads to longer hospitalization.
\nThis pathology occurs due to cytotoxic effects of anticancer drugs/radiotherapy that cause damage at the DNA of stem cell (epithelial cell progenitors) with intense oxidative stress and consequent cell death. This apoptotic process is exacerbated affecting the absorption by shortening the villi structure of enterocytes and causing the loss of epithelial barrier with an invasion of inflammatory cells (neutrophils, eosinophils, and macrophages) leading to an increased production of inflammatory mediators at the mucosal area with consequent epithelial erosion and ulceration. The progressive destruction of mucosal integrity causes the rupture of the tight junctions proteins, leading to an increase in the intestinal permeability with subsequent penetration of commensal microbiota to the submucosal layer generating bacteria translocation which exacerbates the inflammatory process and intensifies the symptoms [53, 54, 55, 56, 57]. Besides, the intestinal microbiota composition is also modified by the chemotherapeutic drugs and radiotherapy action [54, 58, 59] resulting in dysbiosis. After the end of treatment, recovery and restoration of the GIT structure occur [60].
\nBesides IBD and mucositis, it has been reported that intestinal microbiota has an intrinsic effect on metabolism, potentially contributing to several features of the pathophysiology of metabolic syndrome [61, 62]. The metabolic syndrome is an accumulation of various risk factors (glucose intolerance, hyperinsulinemia, hypertension, as well as dyslipidemia) which can often be associated with insulin resistance, hypertension with abdominal fat accumulation, and obesity [63, 64, 65].
\nThe etiology of metabolic syndrome is not well-defined; however there are evident characteristics and life habits that could contribute to its development such as unbalanced diet, smoking, lack of physical activity, and the genetic predisposition [66]. These factors directly increase the risk of cardiovascular disease and chronic diseases as type 2 diabetes mellitus and obesity, and the interaction between components of both the clinical and biological phenotypes of the syndrome contributes to the development of a pro-inflammatory state [67].
\nThe inflammatory process observed in MS is directly associated with increased oxidative stress. The reactive oxygen species (ROS) are capable of mediating symptoms of diabetes mellitus, such as insulin resistance and decrease in insulin secretion, and attend as precursors for the formation of LDLox (oxidized low-density lipoproteins), responsible for a large part of the development of atherosclerotic lesions, and the increase in circulating cholesterol fractions and glucose [68, 69]. In addition, chronic diseases are directly related to changes in the intestinal microbiome [70, 71], and they are also associated with elevated circulating levels of pro-inflammatory cytokines such as TNF and IL-6 [72].
\nThe probiotic use in attenuating symptoms of different inflammatory diseases is widely reported in the literature. Among the commercial probiotics studied for treatment of these diseases, only a few products have been extensively tested in clinical trials in patients with MS, in order to demonstrate an effective result on weight loss, lipid metabolism, and reduction of inflammatory markers.
\nStudies performed with Lactobacillus strains have shown the ability of these probiotics in reducing the lipid accumulation in adipose tissues, as well as in inducing the subexpression of lipogenic genes [73, 74]. Animals that received diets with high concentrations of lipids and then treated with L. gasseri SBT2050 had shown lower intestinal permeability and bacterial translocation, as well as reduction of inflammatory parameters, suggesting that this strain improves the intestinal barrier function [75, 76, 77, 78]. In addition, L gasseri BRN17 was studied to treat animals with MS caused by the carbohydrate-rich diet consumption. This strain reduced the accumulation of adipose tissue in mice, and it has a beneficial effect on weight loss [79, 80, 81]. Another important approach with associated probiotics (Bifidobacterium, Lactobacillus, and S. thermophilus) for treatment of overweight patients has shown an improvement in lipid profile, as well as insulin sensitivity [82]. Besides, recently Hsieh e collaborators [83] demonstrated that administration of live Lactobacillus reuteri ADR-1 and killed Lactobacillus reuteri ADR-3 strain ameliorated type 2 diabetes mellitus in a clinical trial. The results indicated that the consumption of ADR-1 displayed a reduction effect on serum glycated hemoglobin (HbA1c), triglyceride, and cholesterol levels. On the other hand, the intake of ADR-3 showed a beneficial effect on blood pressure reduction. Besides, a reduction in the levels of pro-inflammatory cytokines (IL-1β), increase in antioxidant enzyme (superoxide dismutase), and the changes in intestinal microflora composition (increase in intestinal level of Lactobacillus spp. and Bifidobacterium spp. and decrease in Bacteroidetes) were observed. Thus, these strategies highlight the beneficial and potential effect of interventions targeting gut microbiota modulation by the use of probiotic strains to treat components or complications of metabolic syndrome.
\nThe human being for more than 4000 years has been consuming fermented products, by the fermentation process. At the beginning this practice was done to preserve foods from either physical, chemical, or microbial alterations. The microorganisms participating in this process are the lactic acid bacteria, extensively widespread in nature and also belong to the GIT communities, able to convert the sugar in lactic acid as well as produce other metabolites which contribute to food modifications, either sensorial or nutritional value. Thus, the terminology “functional food” was attributed to food with health benefits to the consumer including nutritional and physiological function [84, 85, 86].
\nDuring the fermentation, these bacteria can contribute to improving the digestion of nutrients (lactose, proteins, small peptides, and polysaccharides); providing essential micronutrients (vitamins) as well as bioactive compounds (metabolites) with potential health benefits to the host, such as prevention against enteric inflammation [87, 88]; providing antimicrobial, antihypertensive, hypocholesterolemic, immunomodulatory, antioxidant, and anticancer effects [46, 85, 89, 90, 91, 92]; showing ability to regulate the immunity; and, consequently, improving host quality of life [93].
\nTherefore, the gut communities and the microbial-derived molecules present in the gut lumen have been strongly influenced, either qualitatively or quantitatively, by consumption of dairy products [94] such as yogurts, cheeses, and fermented milk, among other fermented products using probiotic bacteria. Thus, the microbiota manipulation by functional food, probiotics, and prebiotics are evaluated as a beneficial option for treatment of GIT diseases [95].
\nThere is a constant interaction between the host and the bowel commensal bacterial community in order to maintain the homeostasis [3, 96, 97, 98]. However, when this mutualist relationship is compromised, the intestinal microbiota may cause and/or contribute to either the establishment or the progression of inflammatory diseases [96, 97, 98, 99]. In this context, the search for therapeutic strategies that minimize the development and progression of pathologies caused directly and indirectly by the unbalance of the commensal microbiota has grown. The consumption of probiotic bacteria is one of these strategies, as they present several effects, such as ability to improve the intestinal barrier, stimulate the systemic and mucosal immune system, regulate the composition of the intestinal microbiota, and provide essential micronutrients (such as vitamins and SCFAs) and other bioactive compounds (metabolites) with potential health benefits for the host [100, 101, 102, 103].
\nProbiotics are defined as “live microorganisms that offer host health benefits when administered in adequate amounts” [104, 105]. The majority of the studied probiotics belongs to the group of lactic acid bacteria. However, other microorganisms with probiotic properties also deserve attention, such as yeasts (Saccharomyces spp.) and bacteria of the genus Bifidobacterium and Faecalibacterium, among others [106, 107, 108].
\nLAB, which include, mainly, species from the genus Lactobacillus, Leuconostoc, Lactococcus, Pediococcus, and Streptococcus, constitute a group of Gram-positive, anaerobic or aerotolerant, nonspore-forming, nonmobile, and highly low pH-tolerant microorganisms. However, the main characteristic of this group is its ability to produce lactic acid as the final product of the fermentation of carbohydrates [109, 110, 111].
\nLAB are often present in the human gut but also can be introduced by the ingestion of fermented foods, such as yogurt and other fermented milk products and fermented cured meat by-products [103], having the generally recognized as safe (GRAS) status by the Food and Drug Administration (FDA). Lactobacillus spp., Streptococcus spp., and Lactococcus spp. are the major LAB species with probiotic effects, and they have been used in therapeutic applications for treatment and prevention of various intestinal disorders [112, 113].
\nScientific evidence reveals that the mechanisms by which probiotic bacteria ameliorate inflammatory bowel damage are heterogeneous, strain specific, and dependent on the number of available bacteria. Thus, administration of probiotic bacteria, specially LAB, improves intestinal inflammatory responses by (i) modulation and normalization of perturbed intestinal microbial communities; (ii) competitive exclusion of pathogens such as Staphylococcus aureus and Salmonella typhimurium, among others; (iii) bacteriocin and SCFA production; (iv) enzymatic activities related to metabolization of a number of carcinogens and other toxic substances; (v) adhesion to mucosal cells, cell antagonism, and mucin production; (vi) intestinal permeability reduction by tight junctions protein modulation (e.g., zonulin, claudin, occludin, junctional adhesion molecule); (vii) modulation of the immune system by stimulating Tregs cells, IgA production by B cells, and NF-kβ signaling pathway inhibition; and (viii) interaction with the brain-gut axis via the generation of bacterial metabolites (\nFigure 1\n) [103, 114, 115, 116, 117, 118].
\nA schematic diagram about potential action mechanisms of probiotic bacteria.
In order to potentialize the beneficial effects of probiotic strains, research has been conducted over the last decades, based on genetic engineering techniques, especially those related to DNA manipulation. Thus, modern methods of genetic engineering open the new opportunities to design and create genetically modified probiotic strains with the desired characteristics or to exclusively target a specific pathogen or toxin to be used either as a vaccine or for drug delivery [119, 120]. Since most of the probiotic strains are part of the LAB group, most of the genetic manipulation studies are carried out with species that belong to this group, such as Lactococcus and Lactobacillus genera. Consequently, recombinant probiotics have been created for mucosal delivery of therapeutic and/or prophylactic molecules comprising DNA, peptides, single-chain variable fragments, cytokines, enzymes, and allergens [121, 122], leading to the concept of “biodrug” for the prevention and treatment of various diseases [123]. Thus, researches have emphasized the use of species of these genera in two different approaches: the first as producers of heterologous protein and the second as vehicle for delivery of DNA vaccines [124].
\nMany studies are carried out with Lactococcus lactis due to its economic importance in the production of cheese and its easy growth and manipulation. In addition, it was the first species of LAB to have its genome completely sequenced, which allowed a greater understanding of its genetic and physiological mechanisms, aiding in the development of technological packages for its genetic manipulation in a laboratory environment [124, 125, 126, 127, 128].
\nThere are several ways to make LAB produce heterologous proteins, and the most used form is through the insertion of a plasmid into its cytoplasm. Plasmids are elements of extrachromosomal DNA that are naturally found in prokaryotes. With the advent of the recombinant DNA technique, these elements have been manipulated to act as molecular vehicles that allow the production of proteins of interest by the bacterium [129].
\nThe first heterologous protein production system based on plasmid insertion in LAB was developed for L. lactis. These systems included both inducible and constitutive promoters, which ensure efficient expression of the antigen of interest under different conditions [130, 131]. Although it is possible to choose the type of promoter to be used in the vector, the vast majority of expression vectors present inducible promoters that allow controlled expression of the protein of interest by protecting against aggregation and protein degradation in the bacterial cytoplasm. On the other hand, these vectors present safety issues that need to be analyzed since it is necessary to introduce chemical compounds into the culture medium to induce protein expression prior to animal administration [132, 133, 134].
\nWith the improvement of cloning and expression techniques, several production systems were developed, specifically for LAB, allowing the production of different molecules of interest, including pathogen antigens, by a large number of LAB species [135, 136, 137, 138, 139]. The most commonly used regulation systems in LAB are the following:
\nAmong the heterologous production systems, the most widely studied is the nisin-controlled gene expression system. This system is based on the expression of three genes (nisA, nisF, and nisR) that are involved in the production and regulation of the antimicrobial peptide nisin, which is naturally secreted by different strains of L. lactis. In this system the membrane-located histidine kinase NisK senses the signal inducer nisin and autophosphorylates and then transfers the phosphorous group to the intracellular response regulator protein NisR which acts as a transcription activator of nisA/nisF and induces gene expression under pNis promoter. Depending on the presence or absence of the corresponding targeting signals, the protein is either expressed into the cytoplasm or the cell envelope or secreted into the external medium [140]. Thus, it has already been successfully used for the expression of different proteins of medical and biotechnological interest [141, 142].
\nIn 2004, Miyoshi and colleagues [143] developed the xylose-inducible expression system whose promoter is the xylose permease gene (pxylT) found in L. lactis NCDO2118. This system produces either cytoplasmic or secreted proteins being activated in the presence of xylose and strongly repressed in the presence of glucose, fructose, or mannose [143].
\nMore recently, the stress-inducible controlled expression system was developed using the L. lactis groESL promoter [134]. This system induces expression of proteins of interest via stress stimuli such as those found in the GIT (e.g., bile salt, acid pH, antimicrobial peptide, and heat shock proteins) [134, 144]. This system does not require the induction of bacterial culture or the presence of regulatory genes, being a good alternative in the delivery and production of therapeutic proteins at mucosal surfaces.
\nAmong the available approaches to stimulate efficient mucosal responses, the use of bacterial system for DNA delivery and its expression using the eukaryotic cell machinery have been extensively explored. Unlike the production of heterologous protein, in which the bacterium is responsible for the synthesis of the protein of interest, in the DNA vaccine platform, the bacteria only act as a delivery vehicle for prophylactic and therapeutic purposes [109, 145].
\nNew vectors had been developed to approach the DNA vaccine using LAB as live delivery vehicles [146, 147, 148, 149, 150]. These vectors present a series of common characteristics such as the presence of a eukaryotic promoter, which allows protein expression by eukaryotic cells; a prokaryotic region, which has a selection marker (usually antibiotic resistance); a multiple cloning site, where the open reading frame (ORF) of interest will be inserted; and a prokaryotic origin of replication, which ensures that the plasmid replicates only in prokaryotic cells [151]. Some molecules (IL-10, IL-4, and HSP65) have been cloned in these vectors to evaluate their effect, especially as a treatment approach in diseases related to the bowel [152, 153], as well as reporters (GFP and Cherry) which allowed the understanding of this platform in the mammalian body [148, 154]. Although further studies need to be conducted in order to elucidate whether the cloning of ORFs of interest in these vectors is really effective pointing to disease prevention and treatment, this approach is undoubtedly an important tool for the development of new techniques with potential in the medical clinic.
\nAmong the different techniques used to construct recombinant LAB strains, the most recent is associated with the use of the clustered regularly interspaced short palindromic repeats (CRISPR)-Cas system, based on the use of a system present in several bacterial strains that works as part of the adaptive immune system of bacteria and archaea against the presence of external DNA, such as plasmids and bacteriophages [155, 156, 157, 158, 159].
\nAlthough this system has been studied for more than 30 years [160], it was only in 2013 that the first experiments were carried out emphasizing its use as a tool for genome editing [161, 162]. Evaluating the CRISPR databases, it is possible to observe that about 46% of all bacterial genomes presents the CRISPR-Cas system, and this percentage reaches approximately 63% of the sequenced Lactobacillus genomes [163]. The natural presence of this system in most of the LAB strains expands the possibilities of genetic manipulation of microorganisms of this group, including probiotic ones [164].
\nThe first gene editing experiment in LAB based on the CRISPR-Cas system was conducted by Oh and van Pijkeren [165] where they were able to edit three different regions of the genome, with efficiency up to 100% in the selected clones. After this pioneering work, few others were published focusing on LAB gene editing [166, 167, 168].
\nTherefore, the use of this technology is presented as a widely viable strategy to be applied in LAB, enabling the development of food-grade recombinant strains in order to allow their future use in the clinic [169].
\nThe use of recombinant L. lactis strains, as well as others recombinant LAB strains, using different systems has shown promising results in many studies as an alternative therapy to treat, especially, GIT inflammation and other diseases (\nTable 1\n).
\nMicroorganism | \nGene | \nExpression System | \nInflamation Condition | \nAnti-Inflamatory Properties | \nReferences | \n
---|---|---|---|---|---|
\nL. lactis MG1363 | \nMouse IL-10 | \nSICE | \nMouse model of DNBS-induced colitis | \nRestoration of intestinal architecture; IgA production and IL-6 reduction; Reduced tissue damage | \n[134] | \n
\nL. lactis MG1363 | \nMouse IL-10 and IL-4 | \npValac vector | \nMouse model of DSS/TNBS-induced colitis | \nDecreased IL-6, IL-12 and MPO activity Reduced tissue damage | \n[152, 153] | \n
\nL. lactis NZ9000 | \nMouse TGF-β1; IL-10 and leukocyte protease inhibitor Human Elafin | \nNICE | \nMouse model of DSS-induced colitis | \nReduced tissue damage Decreased pro-inflammatory cytokines | \n[174] | \n
\nL. lactis NCDO 2118 | \nHuman 15-lipoxygenase-1 | \nXIES | \nMouse model of DSS-induced colitis | \nReduced tissue damage | \n[175] | \n
\nL. lactis NCDO 2118 | \n\nM. leprae Hsp65 protein | \nXIES | \nMouse model of DSS-induced colitis | \nRestoration of intestinal architecture CD4+Foxp3+ and CD4+LAP+ regulatory T cells production | \n[176] | \n
\nB. bifidum BS42 | \nMouse IL-10 | \nBEST | \nMouse model of DNBS-induced colitis | \nReduced tissue damage | \n[177] | \n
\nL. casei BL23 | \nSuperoxide dismutase A from L.lactis MG1363 Catalase from L.plantarum ATCC | \npLEM415 vector | \nMouse model of TNBS-induced Crohn’s disease | \nReduced tissue damage Reduced microbial translocation Increase IL-10/INF-γ reduction | \n[180] | \n
\nS. thermophilus CLR807 | \nSuperoxide dismutase A from L.lactis MG1363 Catalase from L.plantarum ATCC | \npIL253 vector | \nMouse model of TNBS-induced colitis | \nReduced tissue damage Reduced microbial translocation IL-17 reduction | \n[181] | \n
\nL.lactis AG013 | \nHuman Trefoil Factor 1 (Htff-1) | \nThyA native promoter of L.lactis\n | \nHamster model of radiation-induced oral mucositis | \nReduced clicnical scores of oral mucositis | \n[186] | \n
\nL. lactis NZ9000 | \nHuman pancreatitis associated protein (Reg3A) | \nNICE | \nMouse model of 5-FU-induced intestinal mucositis | \nMicrobiota Regulation Villus architecture preservation Increased Paneth cells activity | \n[185, 187] | \n
\nL.lactis NCDO2118 | \n\nM. leprae Hsp65 protein | \nXIES | \nMice model of experimental encephalomyelitis | \nIncreased CD4+Foxp3+ regulatory T cells Reduced encephalytogenic CD4+ T cells | \n[184] | \n
\nL.lactis MG1363 | \nMouse IL-17 | \nSICE | \nMice model HPV-induced cancer | \nReduced tumor size Induced IL-6 and IL-17 secretion | \n[182] | \n
\nL.lactis NZ9000 | \n\nM. leprae Hsp65 protein and peptide derived of human Hsp60 protein | \nNICE | \nMice model of diabetes type 1 | \nReduction of insulitis Inhibition of T cell proliferation | \n[183] | \n
Protein with anti-inflammatory properties produced in different strains of bacteria.
To arrive at mucosa in sufficient quantities to exert their therapeutic effects, many LAB strains must survive, during their passage through the GIT, stressor factors such as pH, temperature, bile salt concentration, and the presence of antimicrobial peptides [170, 171, 172]. In this context, an interest approach was recently developed by Coelho-Rocha and colleagues [154] using an encapsulated recombinant strain (L. lactis pExu:mcherry) and tested it through the GIT at different times post-administration. They have shown that the microencapsulation process is an effective method to improve DNA delivery, guaranteeing a greater number of viable bacteria able to reach different sections of the bowel [154].
\nThe use of recombinant probiotics to improve therapeutic approaches has been widely studied using different systems with different molecules. As IBDs are a serious clinical topic, many strategies have been tested trying to improve previous results found with wild type strains.
\n\nL. lactis MG1363 strain carrying the pTREX1 vector expressing the mouse IL-27 protected mice against the inflammatory effects of dextran sulfate sodium (DSS)-induced colitis. This recombinant strain was able to reduce disease activity scores and pathology features of the large and small bowels and also led to reduced levels of inflammatory cytokines IL-1β, TNF-α, and IFN-γ in colonic tissue. In addition, reduction in the number of CD4+ and IL-17+ T cells in gut-associated lymphoid tissue and increase in IL-10 production were observed [173].
\nBesides, it was also demonstrated in a DSS-induced colitis mouse model that the oral administration of L. lactis NZ900 strain harboring the NICE system expressing either the anti-inflammatory cytokine IL-10, TGF-β1, secretory leukocyte protease inhibitor (SLPI), or elafin was able to ameliorate some clinical parameters in inflamed mice. Even though it was possible to observe the reduction of weight loss and diarrhea, microscopic colonic damage scores, colon thickness, and myeloperoxidase (MPO) activity, the authors reported that treatments with recombinant L. lactis strain delivering either SLPI or elafin were more efficient to reduce signs of colitis than treatments with anti-inflammatory cytokines. Altogether these recombinant strains display anti-inflammatory effects in inflamed mice [174].
\nApproaches using the invasive L. lactis MG1363 FnBPA+, by expressing the FnBpA protein at their surface and carrying the pValac eukaryotic expression vector coding either the IL-10 cytokine [rL. lactis FnPBA+ (pValac:il-10)] or the IL-4 cytokine [rL. lactis FnPBA+ (pValac:il-4)] in DSS or trinitrobenzenesulfonic acid (TNBS)-induced acute model of colitis, respectively, were also investigated. The administration of L. lactis FnPBA+ (pValac:il-10) recombinant strain was capable to reduce the intestinal inflammation by increasing IL-10 levels and sIgA production, accompanied by decreasing IL-6, as well as the restoration of intestinal architecture of mice colon [153]. Besides, the engineered L. lactis FnPBA+ (pValac:il-4) was able to slump the level of pro-inflammatory cytokine (IL-12, IL-6) and myeloperoxidase activity and increase levels of IL-4 and IL-10, consequently decreasing the colitis harshness [153].
\nThe human 15-lipoxygenase-1-producing L. lactis NCDO2118 harboring the xylose-inducible expression system (pXylt:CYT:15-LOX-1) was also effective in attenuating the symptoms of DSS-induced colitis in a murine model [175]. Its oral administration improved the body weight, decreased pro-inflammatory cytokines (IFN-γ and IL-4) while increasing the anti-inflammatory cytokine IL-10, and, consequently, ameliorated the macroscopic damage scores associated with the inflammation.
\nThe oral pretreatment with genetically modified L. lactis NCDO2118 able to secrete HSP65 protein from Mycobacterium leprae, using XIES system (pXylt:SEC:hsp65), prevented DSS-induced colitis in C57BL/6 mice [176]. This protection was associated with reduced pro-inflammatory cytokines, such as IFN-γ, IL-6, and TNF-α; it also increased IL-10 production in colonic tissue and expansion of CD4+FoxP3+ and CD4+ latency-associated peptide (LAP+) regulatory T cells in the spleen and mesenteric lymph nodes. Besides, the authors showed that this effect was dependent on IL-10 and toll-like receptor 2 (TLR-2) [176].
\nAlthough L. lactis represents an excellent candidate for a live mucosal vector delivery system, other bacteria have also been explored as promising live vehicles for molecule expression with therapeutic properties, such as Lactobacillus, Bifidobacterium, and Streptococcus. In this context, Mauras et al. [177] using the new Bifidobacteria Expression SysTem (BEST) allowing the production of IL-10 in Bifidobacterium bifidum BS42(pBESTExp4:il-10 and pBESTBL1181:il-10) demonstrated that the use of these recombinant strains in a DNBS-induced colitis model showed its ability to decrease local inflammation and confirmed therefore its potential for delivery of therapeutic molecules in the colon.
\nIt is well known that IBD is associated with oxidative stress by the increase in concentration of reactive oxygen species in the GIT and impaired antioxidant defenses [178, 179]. In this context, it has been shown that some probiotic LAB strains may play a protective role in IBD by expressing antioxidant enzymes such as superoxide dismutase (SOD) and catalase (CAT) [180, 181].
\nLeBlanc et al. and Del Carmen et al. [180, 181] showed, respectively, that L. casei BL23 and S. thermophilus CRL807 transformed with two different plasmids (pLEM415:mnkat; pLEM415:sodA) (pIL253:sodA and pIL253:mnkat) harboring the genes encoding catalase (CAT) or superoxide dismutase (SOD) antioxidant enzymes exhibited anti-inflammatory activities in a mouse model of Crohn’s and colitis disease induced by trinitrobenzenesulfonic acid (TNBS). The authors observed a reduction in weight loss, fewer liver microbial translocation, lower macroscopic and microscopic damage scores, and modulation of the IFN-γ/IL-10 [180] and IL-10/IL-17 [181] cytokine production in the large intestines of mice treated with either CAT- or SOD-producing lactobacilli/streptococci.
\nThe stress-inducible controlled expression (SICE) system represented by L. lactis MG1363 strain harboring the pLB333 plasmid was developed to avoid the external induction of culture before the host administration [134]. Several interesting molecules were cloned in this system such as IL-10 [134] and IL-17 [182], and the effect of L. lactis secreting them was evaluated in mice models. L. lactis (pSICE:il-10) was tested in a DNBS-induced colitis mice model, resulting in a significant reduction in colitis parameters with improvement in weight loss and a decrease in macroscopic scores [134]. The intranasal administration with L. lactis secreting IL-17A (pSICE:il-17), in a mice model of human papilloma virus (HPV)-induced cancer, was able to reduce tumor size and induce IL-6 and IL-17 secretion in reactivated splenocytes from mice challenged with the tumoral cell line [182]. Both works confirmed the potential use of L. lactis harboring the SICE system to deliver interesting molecules either to colitis or colon cancer patients [134, 182].
\nAlthough many studies have focused on the use of recombinant bacteria for the treatment of IBDs, as was previously discussed, the use of recombinant probiotic strains expressing/delivering therapeutic molecules has been explored for treatment or prevention of other diseases such as mucositis, cancer, obesity, multiple sclerosis, and diabetes [182, 183, 184, 185].
\nAn in vivo study reported by Caluwaerts et al. [186] showed that recombinant L. lactis AG013 secreting human trefoil factor 1(hTFF-1) was able to reduce the severity and course of radiation-induced oral mucositis. Carvalho et al. [187] also demonstrated that a recombinant strain of L. lactis NZ9000 using the inducible NICE system to express the human pancreatitis-associated protein (PAP) was able to prevent 5-FU-induced intestinal mucositis in a murine model. It was observed that this protein preserved villous architecture, increased Paneth cell activity [187], and suppressed the growth of Enterobacteriaceae during inflammation [185].
\nIt also has been shown that oral administration of a recombinant L. lactis NCDO2118 strain (pXylT:SEC:hsp65) prevented the development of experimental autoimmune encephalomyelitis (EAE) in C57BL/6 mice [184]. Mice fed daily with this recombinant strain increased the number of natural and inducible CD4+FoxP3+ and CD4+ latency-associated peptide (LAP+) regulatory T cells in the spleen, inguinal and mesenteric lymph nodes, as well as in the spinal cord. In addition, a reduction in the recruitment of encephalitogenic CD4+ T cells to the spinal cord was observed, which decreased IgG response against HSP65 and induced an anti-inflammatory cytokine profile (IL-17 reduction and IL-10 increase) during EAE development.
\nThe oral administration of recombinant L. lactis expressing HSP65 and tandemly repeated P277 (pCYT:HSP65-6P277) was also analyzed in a model of type 1 diabetes mellitus (DM1) [183]. The authors observed that oral administration of recombinant L. Lactis resulted in the prevention of hyperglycemia, improved glucose tolerance and reduced insulitis, and induced HSP65- and P277-specific T-cell immunotolerance, as well as antigen-specific proliferation of splenocytes, demonstrating to be an effective therapeutic approach in preventing DM1 [183].
\nAnother study using the E. coli Nissle 1917 strain engineered to secrete N-acylphosphatidylethanolamines (NAPEs) (pDEST-At1g78690 expression plasmid) demonstrated that this strain was able to reduce the obesity of mice fed with a high-fat diet when added to drinking water. N-acyl phosphatidylethanolamines are precursors to the N-acylethanolamine (NAE) family of lipids, which are synthesized in the small intestine in response to feeding and reducing food intake and obesity. Mice that received modified bacteria had dramatically lower food intake, adiposity, insulin resistance, and hepatosteatosis than mice receiving standard water or control bacteria [188]. In addition, it was observed that changes on intestinal microbiota significantly decreased the abundance of Firmicutes and increased the abundance of Proteobacteria. Thus, these results provide evidence of the potential efficacy of this approach to inhibit the development of metabolic disorders and related diseases.
\nCurrently the association between disease progression, especially chronic inflammatory diseases, and intestinal dysbiosis has been more frequently observed. As a clinical strategy, the use of probiotic bacteria, which naturally benefit the host, has been increasingly used on the treatment of diseases related to the GIT. In view of the good results obtained with this approach, researchers have sought through bacterial genetic modification to increase the beneficial potential of probiotics, either through their use for heterologous protein production or as a vehicle for vaccinal plasmid delivery, by developing recombinant bacterial strains and by testing their action in different disease models. And while there are still a number of questions that need to be answered about the use of genetically modified organisms for health care, especially in human, the use of these strains has proven to be a potentially effective therapeutic alternative, so much so that clinical trials using recombinant lineages have already been authorized and conducted in humans.
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I am also a member of the team in charge for the supervision of Ph.D. students in the fields of development of silicon based planar waveguide sensor devices, study of inelastic electron tunnelling in planar tunnelling nanostructures for sensing applications and development of organotellurium(IV) compounds for semiconductor applications. I am a specialist in data analysis techniques and nanosurface structure. 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After obtaining a Master's degree in Mechanical Engineering, he continued his PhD studies in Robotics at the Vienna University of Technology. Here he worked as a robotic researcher with the university's Intelligent Manufacturing Systems Group as well as a guest researcher at various European universities, including the Swiss Federal Institute of Technology Lausanne (EPFL). During this time he published more than 20 scientific papers, gave presentations, served as a reviewer for major robotic journals and conferences and most importantly he co-founded and built the International Journal of Advanced Robotic Systems- world's first Open Access journal in the field of robotics. Starting this journal was a pivotal point in his career, since it was a pathway to founding IntechOpen - Open Access publisher focused on addressing academic researchers needs. Alex is a personification of IntechOpen key values being trusted, open and entrepreneurial. Today his focus is on defining the growth and development strategy for the company.",institutionString:null,institution:{name:"TU Wien",country:{name:"Austria"}}},{id:"19816",title:"Prof.",name:"Alexander",middleName:null,surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/19816/images/1607_n.jpg",biography:"Alexander I. Kokorin: born: 1947, Moscow; DSc., PhD; Principal Research Fellow (Research Professor) of Department of Kinetics and Catalysis, N. Semenov Institute of Chemical Physics, Russian Academy of Sciences, Moscow.\r\nArea of research interests: physical chemistry of complex-organized molecular and nanosized systems, including polymer-metal complexes; the surface of doped oxide semiconductors. He is an expert in structural, absorptive, catalytic and photocatalytic properties, in structural organization and dynamic features of ionic liquids, in magnetic interactions between paramagnetic centers. The author or co-author of 3 books, over 200 articles and reviews in scientific journals and books. He is an actual member of the International EPR/ESR Society, European Society on Quantum Solar Energy Conversion, Moscow House of Scientists, of the Board of Moscow Physical Society.",institutionString:null,institution:{name:"Semenov Institute of Chemical Physics",country:{name:"Russia"}}},{id:"62389",title:"PhD.",name:"Ali Demir",middleName:null,surname:"Sezer",slug:"ali-demir-sezer",fullName:"Ali Demir Sezer",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/62389/images/3413_n.jpg",biography:"Dr. Ali Demir Sezer has a Ph.D. from Pharmaceutical Biotechnology at the Faculty of Pharmacy, University of Marmara (Turkey). 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Focus of his research activity is drug delivery, physico-chemical characterization and biological evaluation of biopolymers micro and nanoparticles as modified drug delivery system, and colloidal drug carriers (liposomes, nanoparticles etc.).",institutionString:null,institution:{name:"Marmara University",country:{name:"Turkey"}}},{id:"61051",title:"Prof.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"100762",title:"Prof.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"St David's Medical Center",country:{name:"United States of America"}}},{id:"107416",title:"Dr.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Texas Cardiac Arrhythmia",country:{name:"United States of America"}}},{id:"64434",title:"Dr.",name:"Angkoon",middleName:null,surname:"Phinyomark",slug:"angkoon-phinyomark",fullName:"Angkoon Phinyomark",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/64434/images/2619_n.jpg",biography:"My name is Angkoon Phinyomark. I received a B.Eng. degree in Computer Engineering with First Class Honors in 2008 from Prince of Songkla University, Songkhla, Thailand, where I received a Ph.D. degree in Electrical Engineering. My research interests are primarily in the area of biomedical signal processing and classification notably EMG (electromyography signal), EOG (electrooculography signal), and EEG (electroencephalography signal), image analysis notably breast cancer analysis and optical coherence tomography, and rehabilitation engineering. I became a student member of IEEE in 2008. During October 2011-March 2012, I had worked at School of Computer Science and Electronic Engineering, University of Essex, Colchester, Essex, United Kingdom. In addition, during a B.Eng. I had been a visiting research student at Faculty of Computer Science, University of Murcia, Murcia, Spain for three months.\n\nI have published over 40 papers during 5 years in refereed journals, books, and conference proceedings in the areas of electro-physiological signals processing and classification, notably EMG and EOG signals, fractal analysis, wavelet analysis, texture analysis, feature extraction and machine learning algorithms, and assistive and rehabilitative devices. I have several computer programming language certificates, i.e. Sun Certified Programmer for the Java 2 Platform 1.4 (SCJP), Microsoft Certified Professional Developer, Web Developer (MCPD), Microsoft Certified Technology Specialist, .NET Framework 2.0 Web (MCTS). I am a Reviewer for several refereed journals and international conferences, such as IEEE Transactions on Biomedical Engineering, IEEE Transactions on Industrial Electronics, Optic Letters, Measurement Science Review, and also a member of the International Advisory Committee for 2012 IEEE Business Engineering and Industrial Applications and 2012 IEEE Symposium on Business, Engineering and Industrial Applications.",institutionString:null,institution:{name:"Joseph Fourier University",country:{name:"France"}}},{id:"55578",title:"Dr.",name:"Antonio",middleName:null,surname:"Jurado-Navas",slug:"antonio-jurado-navas",fullName:"Antonio Jurado-Navas",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/55578/images/4574_n.png",biography:"Antonio Jurado-Navas received the M.S. degree (2002) and the Ph.D. degree (2009) in Telecommunication Engineering, both from the University of Málaga (Spain). He first worked as a consultant at Vodafone-Spain. From 2004 to 2011, he was a Research Assistant with the Communications Engineering Department at the University of Málaga. In 2011, he became an Assistant Professor in the same department. From 2012 to 2015, he was with Ericsson Spain, where he was working on geo-location\ntools for third generation mobile networks. Since 2015, he is a Marie-Curie fellow at the Denmark Technical University. 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