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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:"72",title:"Ionic Liquids",subtitle:"Theory, Properties, New Approaches",isOpenForSubmission:!1,hash:"d94ffa3cfa10505e3b1d676d46fcd3f5",slug:"ionic-liquids-theory-properties-new-approaches",bookSignature:"Alexander Kokorin",coverURL:"https://cdn.intechopen.com/books/images_new/72.jpg",editedByType:"Edited by",editors:[{id:"19816",title:"Prof.",name:"Alexander",surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"64961",title:"Doing and Being: A Metaphysic of Persons from an Ontology of Action",doi:"10.5772/intechopen.82837",slug:"doing-and-being-a-metaphysic-of-persons-from-an-ontology-of-action",body:'A significant and worrying lacuna lies at the heart of neuroethical debate. What it lacks is the anchor of a
This chapter aims to supply that anchor, to articulate a conception of persons that will overcome this piercingly divisive dichotomy. It does so, not by privileging one side over the other; a pointless exercise in any event, since neither one is coherent and, besides, they terminate in the coinciding of reductive and flattened abstractions with inflationary, transcendentalised ones. Rather, the dichotomy is overcome by a conception of consciousness grounded in action: action as essentially personal; actions owned, intended; actions that embody those intentions, embody
It is in this empirical sense—the philosophically well-brought-up reader may be reassured to learn—that our conception of persons is to be understood as metaphysical. Our aim, in short, is not to critique the neurosciences or rebut their discoveries. It is no part of our case to deny the role played by understanding the brain and brain-function in understanding consciousness and ‘personhood.’ We wish, rather, simply to demonstrate that—if we may be forgiven—there is more to persons than meets the fMRI.
Personal action is ontologically primitive; it is also empirically, which is to say experientially, irrefragable. I cannot deny the reality of my actions without self-stultification, let alone self-contradiction; no more can I deny the actions of others, actions in which my own are but one ingredient. Action is the foundation, the condition, of experience, so meets Ockham’s razor, edge to edge. As such, action is also
Put simply, personal action is an anti-metaphysical metaphysics. As such, it is also an
There is one further point before embarking on the discussion proper. What follows operates solely from a philosophical perspective; for it is this perspective, we are reliably informed, that neuro
Let us begin with an account, in general terms, of the philosophical problem circumscribed by this lacuna in the discourse.
It is tempting, at first, to state the obvious and assert that the dichotomy threatening to tear neuroethics asunder is a product of reductive physicalism or philosophical materialism. Such reductivism is, after all, characteristic of the scientific method that determines the course of neuroscience and so must inform the neuroethicist’s outlook. In consequence, said neuroethicist will inevitably identify consciousness with the neurological, i.e. physical, processes mapped by fMRI scans and, therefore, persons with brains. All this may be true. And yet, we would do well to remember that the obvious does not always stand on solid philosophical ground; besides which, the assertion is easily countered.
During the last century, the physical sciences have seen such extraordinary—one might even say
So much for tilting at windmills. In reality, we surrender to that first temptation and point our superior philosophical finger at the neuroscientist, only to commit the very mistake we accuse her of, thereby reinforcing an already apparently intractable conflict between two different modes of thinking. Fortunately, it is an important principle of our anti-metaphysical metaphysics that pointing fingers point in two directions at once: acts unfold in consequences, so identify the objects on which they bear; simultaneously, however, they reveal intentions and, crucially, the agent of intentions. The root of the problem, that is, lies not in a faulty science, but in bad philosophy; our obvious assertion is itself symptomatic of the very confused and erroneous thinking that gives rise to the problem. We have, in short, transformed method into metaphysic, and a wholly incompatible metaphysic at that.
The results are incompatible because the transformation issues in some form of realism. It is not, perhaps, that naïve realism which supposes, in Russell’s pithy phrase, ‘things are what they seem.’ Nevertheless, it is close cousin to that self-same ‘plebeian illusion’ which Einstein described, ‘according to which things “are” as they are perceived by us through our senses;’ excepting that, in this instance, experiment and observation substitute for sensory perceptions [2, p. 20]. In fact, this substitution means that our method-cum-metaphysic mirrors most closely Peter Byrne’s ‘innocent realism’ [3]. This, we are told, ‘merely reflects on the content of our empirical claims, notes that most of them do not speak about how the world looks from a human perspective and concludes that the world, its things and properties, is for the most part independent of us and our representations’ [3, p. 40]. That we do not articulate our presuppositions, it does not, of course, follow that there are none to articulate. However, the point is clear: no matter what the epistemic medium may be, we are still claiming to identify, to know about, a world that lies, logically and ontologically, beyond the reach of any actual or possible experience.
Any such claim must prove deeply problematic for the empirically minded, not least because it marks an attempt to found scientific knowledge on that which is
Those maps, models, and diagrams are endowed with objectivity by the formulation and application of rules for their construction, rules that constitute the theoretical framework within which any scientific enquiry must be pursued. The more completely and systematically those rules can be defined, the more likely it is they will supply objective facts; but they are not and cannot be ‘independent of us and our representations’ as the realist imagines. They are objective insofar as they overcome the limitations of the individual enquirer’s perspective by abstracting from the subjective immediacy of ordinary sense experience. As the philosopher and physical chemist, Michael Polanyi explained, rules disregard the individual’s ‘normal approach to experience,’ so remain ‘unaffected by the state of the person accepting… [them].’ They come ‘between our senses and the things of which our senses otherwise would have gained a more immediate impression,’ so regulate the organisation and interpretation of those impressions. What is more, and in some ways more important, those rules are open to evaluation by all those qualified and equipped to do so: viz., the community of enquirers. Hence, their objectivity is underwritten by universal acceptance: the acceptance of all those participating in scientific research, whatever their field [5, pp. 3–4].
To suggest, as we have done, that empiricism and metaphysical realism are incompatible may be strictly true, but it is also, in one rather limited sense, somewhat misleading. In fact, there comes a point within the rationalist’s abstract conceptualising when the opportunity arises for, not merely for compatibility, but for full-blown coincidence. This is the precise point at which realism becomes idealism and vice versa.
For realism, the point arrives when it finally acknowledges the implications of its supposedly ‘minimal dualism’: ‘how we say things are is one thing, how things
The root cause of this metaphysical mistake lies in the assumption that the neuroscientist’s models and diagrams obtain a precise correspondence with the objects modelled and diagrammatised. In representing the biochemical processes of the brain, it is supposed, the fMRI scanner supplies a literal image of, not the corollary of consciousness, but of consciousness itself. Persons, then, are at most a product of, and at least equivalent to, those biochemical processes.
The difficulties that beset such reductive conceptualising are both numerous and notorious; not least, is the tendency to eliminate the moral subject, thereby rendering the whole neuroethical debate redundant. There is little profit to be had from arguing about the moral properties and capabilities of physical processes which are incapable of choice and therefore of responsibility. Any attempt to do so can be no more than anthropomorphic projection: the imaginative conception of impersonal forces as personal ones, which are themselves, we must remember, reducible to the very forces being imaginatively conceived. The rank confusion and, indeed, circularity, entailed by such a move is, we trust, entirely obvious.
A more serious problem, however, may be that equating persons to sheer physical process threatens to eliminate the possibility of meaningful discourse. It does so because, in and of themselves, physical processes do not possess logical properties. The firing of neurones may occur or it may not, but such an event cannot be
It is worth repeating, for clarity’s sake, that it is not the reductive materialism, so called, of the neuroscientist or her
Put simply, the neuroscientist may, if she chooses, conceive of the subjects of her research in purely physical terms, but she cannot conceive herself in the same way. Deny this, and she must concede that her own descriptions of neurological phenomena and all the activities that give rise to them are themselves purely physical phenomena. As such, they must be governed by the same laws of cause and effect that govern all other physical phenomena. There can be no exceptions: the formulation of hypotheses, the devising and performing of experiments to test them, the analysing of results; the sharing of ideas: none of these events can be governed by meaning.
But that is absurd. The neuroscientist’s experiments do not occur, either by accident, or as a function of causal impacts; no more than do the institutions in which neuroscientists work. They are intended activities that some
Having styled herself, no doubt unwittingly, after Descartes’
The bridge between consciousness and the world is broken. We no longer have the means to identify other persons or even ourselves, let alone effect any kind of moral impact. The question we must face, then, is this: under such circumstances of Cartesian ego-isolation, what, in the end, is neuroethics actually
To answer that question, we need only return to our empirical starting point. Consciousness must be reconnected with the world; an easy task since we have, the sciences remind us, the very tools to hand. As Ludwig Feuerbach put it, ‘the necessity of this connection is only sensation’ [12, p. 52]. So saying, Feuerbach admonishes us to reject the demand for mind-independent reality and turn instead to those engaged in exploring and explaining the world, those for whom ‘[t]ruth, reality and sensation are identical’ [12, p. 51]. Only there we shall find the conditions of real knowledge. In their activities, he argued, we may plainly see that ‘[o]nly a sensuous being is a true and real being. Only through the senses and not through thought for [or in] itself is an object given in a true sense.’ Crucially, we must be as rigorous as Feuerbach in the application of this principle, so insist that ‘not only the external but also the internal, not only the flesh but also the mind, not only the object but also the ego are objects of the senses’ [12, p. 58].
That ‘sensuousness’ lays the foundations for a more cogent and, ultimately, altogether practical epistemology. At the same time, it provides the terms for constructing—or perhaps more accurately,
While it is perfectly true that, for Feuerbach as for Heidegger, ‘[t]o-be-here [
Metaphysically speaking, it follows from this that consciousness and the world are ontologically co-terminus: the two cannot be separated, are not ‘abstractable in isolation as a subject that has then to be put in relation to an object’ [13, p. 377]. Instead, Hampshire agreed, we are all of us
Otherwise put, the very possibility of self-identification depends logically on being one ‘self-moving body among other bodies’ [6, p. 46]. The ‘being’ that is
All of which means that our Feuerbachian
What we have on our hands, philosophically speaking, is a logically and ontologically primitive conception of human being as physically (and, ultimately therefore, socially) embodied. Embodiment delimits the worldly physical fact that consciousness, personality, is, so determines the self, locating it in one place
No inert substance, then, but the locus of a self-moving, self-directing agency; bodily existence is the focal point from which the impacts and interactions wherein consciousness elaborates and extemporises itself are expressed. Hence, Feuerbach’s avowal: ‘I am a real, sensuous being, and, indeed, the body in its totality is my ego, my essence itself.’ Otherwise put, the body supplies consciousness with that much-needed
If consciousness is to be sufficiently determinate to know anything or do anything, then it must, in Farrer’s phrase, be ‘perfectly embodied;’ at once, both subject and object of experience, consciousness is a feature of the world and so ‘does nothing here without the body’ [8, p. 60]. Crucially, it is this capacity for doing that supplies the ontological and epistemological foundations of a concrete— i.e. combinatorial—ego-profile. That is the ground upon which we shall build our anti-metaphysical metaphysics. Put simply, the physical extensions of consciousness supply our criterion of real being. They do so, because our first and most fundamental experiences are, as the empiricist knows full well, objects ‘of the senses, perception and feeling’ [12, p. 55]. So saying, Feuerbach would use action, more properly
In accessing our environment, so to speak, we are not simply pushing on an open door. We are not, as Farrer put it, ‘swimming in a perfectly featureless medium;’ there is no action
It follows from this that the resistance activity by which our environment is known cannot be random or arbitrary as such; otherwise our attempts to understand and ultimately control it would be fruitless. Without regularity and predictability, consciousness would have no purchase on the world. Hence, as Farrer pointed out, ‘[o]ur conscious experiences find themselves from the start framed by this system’ this regularity and predictability [1, p. 67]. Consequently, experience of resistance, and our engagement with it, take the form of systematic intercourse or controlled interference; that, in turn, supplies consciousness with ‘shape’ or ‘form’. In other words, the interplay between self-activity and resistance activity supplies what Farrer described as the ‘natural unit of thought’ [16, p. 210]. Apart, that is, ‘from my experience of impinging upon, and being impinged upon by, other things or forces, I have no conceivable clue to physical existence, or physical force, or physical interaction’ [16, p. 210]. This is Farrer’s ‘causal solution’ to the problems of realism, ‘minimal dualism’. The world, he reminds us, ‘is not known but as the playground of human thews and human thoughts; were there no free play, there would be no knowledge’ [8, p. 171]. Subject and object are therefore disclosed to one another only as agents of ‘free play’; the features or furniture that occupy our field of action alongside us ‘only become features and so perceptible in so far as they disturb and diversify the field’ [1, p. 234]. In short, the world is (recognisable as) a field of conscious activity and real knowledge is a product of our encounters in and with this field: one may come to know an object ‘only in so far as it varies the disturbances of… [one’s] field—[one] knows it as a class of disturbances.’ We encounter ‘real being’ as it exercises resistance activity; we recognise it by the ‘imprint’ it leaves on our exploratory activities.
To speak ‘objectively,’ then, the world is no more or less than the combination of forces that are continuous with our active explorations. This means that the resistance activities by which the world is known to us and the controlled interference that constitutes our knowing acts are necessarily coeval: consciousness-and-the-world—if we may reiterate a central point—are co-constructed, actualised
This takes us to the ground level of a coherent epistemology. It is also the foundation stone of our metaphysics. To explain: from all that has gone before, it follows that reality as it is known, both by ordinary agents and the most scientifically well-equipped investigators, is to be found, not in inert stuff or substance, but in dynamic process. In Farrer’s Latin phrase,
Action, disturbance-effect, is our metaphysical ultimate;
Thus, to identify what philosophers of mind used to call the ‘seat of consciousness,’ Farrer averred, we need only allow consciousness to pick its own seat by sitting in it [8, p. 24]. Do so, and we shall find that the physical ‘seat’, or more dynamically, ‘vehicle’, of consciousness is located, not in any one phase or feature of the bodily process
This is not, if we may repeat ourselves for a second time, in anyway intended to deny or even diminish the role of the brain in conscious, personal agency. Rather, it is to bestow upon the brain its rightful role and place within the larger, bodily process, which
This is true even when that vehicle does not appear to be moving very much at all. Thinking, for example, about how to frame this sentence is an action and so requires a ‘nerve-plant’ to embody it, however foreshortened the ‘plant’ may be. For thinking, Farrer, reminds us, is the ‘shadow of doing’ and so ‘must be interpreted by a full-blooded doing’ [8, p. 39]. (One suspects that this is the point where many a neuroscientist and neuroethicist commits their fatal error, mistaking this act of interpretation by means of the clue or model of bodily action for ostensive indication or direct denotative reference; and that way, as we have seen, lies metaphysical realism.) For thought apart from overt behaviour, Farrer found an interpretive key right under our noses, so to speak. ‘The best sort of characterisation of thinking is that it is a sort of talking to ourselves’ [7, 29]. The ‘shadow-patterns’ of thought follow same route as speaking, from brain to mouth, taking in lips, jaws, tongue, vocal chords and so on. But they do not get so far: the action-pattern is not fully enacted and the ‘nerve-plant’ fails to flower in ‘full-blooded doing.’ In this way, the act of thinking ‘ghosts’ the act of speaking, stops short of engaging the full physical apparatus of bone and muscle.
The risk of physical reduction here is palpable. As we trace out the route of our action plant, it ill behoves us to ignore the ontological dangers that lie in wait: the abstractions and disjunctions, the dissolution of consciousness into confusion and self-contradiction. Forewarned is forearmed, however; almost literally in this case for, as Farrer pithily put it, we ‘still have mind on our hands just as much as matter’ [8, p. 7].
In fact we have already hinted at the answer more than a little. It lies in the fundamental requirement to make sense of human action as meaningful; to recognise and understand it as governed, not by the diagrammatic laws of cause and effect, but by the rules of discourse and the conventions of the community in which we act. It lies, in short, in the logic of intending. Such logic is essentially presuppositional. It means that the very concept of action in the full and personal sense—the sense, that is, in which we experience it directly in ourselves and the other persons with whom we interact—is only complete when coupled to an intending agent: the owner of the act. In acting, the agent instantiates both the intentional and consequential motifs that make agency what it is: the ‘before’ and ‘after’ of an
Evidently, we have no wish to re-open a logical and ontological chasm so recently closed; equally, no simple reduction will do. Instead, Farrer held out for an agency ‘overplus’ or ‘prior actuality’, insisting that ‘[t]he
We are not, as all of us are no doubt aware, acting and intending in a vacuum. Action and, indeed, everything we have said about it, locates us in a physical
Put simply, acting persons aspire to a lively moral perspicuity by adopting what Charles Conti describes as a ‘metaphysical [i.e. ethically informed] vantage point’ [20, p. 185]. We seek thereby to oversee the means towards realising consequences we actually intend and so avoid colliding with other agents. We view our proceedings, then, not as a ‘Cartesian cogitator but as actor-self and monitor-self simultaneously;’ and so ‘perform our being as we experience it’ [20, p. 185]. In such performances the Cartesian
The social orientation of action coincides—and does occasionally collide—with the ‘internal’ world of conscious deliberation: ‘We sense our compresence with others, so intuit the obligation to act’ [20, p. 186]. Alive to that ‘compresence’, conscious agency is quickened by the possibilities of physical interaction, personal intercourse. That defines the obligation
Unearthing the roots of thought and action, we find that the logic of intending underwrites the concept ‘person’ as a social reality. Logic is not, however, always the most reliable guide to what does and does not exist.7 In view of our much-vaunted empiricism, something a little more concrete would, no doubt, be appreciated. After all, as Farrer reminds us, ‘[i]t is not as though we believed in our neighbour’s personality
Consciousness, then, is awakened, better still
In this way, others supply the terms and conditions of our actions and transactions, thereby staking their claim to the very self they helped create, instilling it with what Feuerbach called ‘the inner life of man’: our social self, our ‘species being’ [15, p. 2]. Like her talk, the other is internalised, metaphysically and morally incorporated into the structure of the self. This process displaces the subjectivity of the subject: its needs, activities, perspective—all felt as intrinsically, immediately present and real,
Taking this one step further, it is, perhaps, sufficiently well known that the derivation of the word ‘person’ lies in the Latin
The essence of consciousness, of ‘personhood’, is fragmentary, consolidated by these exchanged perspectives. This is a commonplace of postmodern identity theories as well as the ‘metaphysical personalism’ (as Conti’s titled his exegesis of Farrer) that we have been mapping here. The ‘unity’ we call a self is actually a function of that primary dialectic of perspectives, the love-relationships into which we are born. In this way, those who had and held us have inexorably bound themselves, their image, into our
It seems we have, at last, reached the philosophical bottom-line. These first and most fertile encounters shape the development of conscious thought and action; they are the grounds which supply form and purpose, sometimes even content, to our explorations and explanations. This is the well-spring of human being, in Martin Buber’s poignant phrase, the ‘cradle of real life’ [22, 29].
Here, then, is an opportune moment to take stock. Let us make the point of moral application plain. Immediately obvious is the absence of any ethical theory, our conception of persons as active agents offers no system or set of rules for the formulation of normative judgements. Being rooted in the personal relations wherein we all, quite literally, find ourselves, our anti-metaphysical,
Applied metaphysics may leave us without a moral theory, but it does not leave us empty-handed. Instead, it supplies the very anchor that our normative judgements demanded from the start: concrete personal connection, the embodiment of moral agency. This rebuts absolutely that Cartesian ghost in—or rather
Philosophically speaking, of course, we have found more than a moral anchor; we also have a coherent logical and ontological framework for our discourse. Personal action supplies the context in which we may clearly see both the particular and the general: first, the analyses and judgements of neuroethics; second, the discipline as a whole and all its participants. Within this framework we may recognise, then understand, and finally overcome the ‘self-sufficing speculation,’ [15, p. xxxiv] which threatens to undermine our efforts. On the one hand, we recall the personal presuppositions of our empiricism: the epistemological requirements of exploring agents that reconnect experience with action, real knowledge with the controlled interference which is the neuroscientist’s stock in trade. On the other, it reveals and resists the temptation to align methodology too closely with metaphysics. This, in turn, allows us to reconcile those binary oppositions—mind and body, intending and intended, subject and object—which do so much to incapacitate every branch of moral philosophy. Reconciliation comes, not by over-inflating empiricism with the transcendental pretensions of metaphysical realism, but by returning us to the only place where those abstract notions can possibly make sense. Mind is a mode of bodily action, body the physical manifestation of mind. Intending and intended are phases of that manifestation, conceptually separable but in reality, i.e. in action, continuous. Subjectivity is essentially other-oriented by virtue of being a reflection of the other who invokes and evokes it in us. Ethically, it denotes obligations owned: my responsibilities as presupposed and, moreover, delimited by my capacity to act in response to a physical and social or personal environment. Being a communal act, objectivity is coeval with this environment: it represents the truth-conditions and epistemic norms laid down by the community of knowing persons. Thus, subject and object are not independent as such, but theoretical perspectives, ways of seeing, of thinking about and understanding the world, the use of which is sanctioned by that community. This does not detract from their truth-value but merely reminds us of the context in which they are first negotiated and defined; that is, transacted with the world by the community of thinkers. Both ethically and epistemologically, then, these theoretical perspectives represent, in their contrasting but congruent ways, the very ‘claimingness’ of others that is our anchor.
Ultimately, then, being firmly anchored by our concept of persons to the solid, social, and inherently ethical ground that entails it, uncouples neuroethical analyses from the arbitrary dictates and philosophical fiats of classical rationalism-cum-realism. Diverted from the rabbit hole of incoherence and irrelevance, which awaits so much philosophical discourse, and possessed of a renewed social conscience, our thoughts and actions are oriented back towards the communities in which even neuroethicists must live and work. Most immediately, perhaps, is the scholarly community whose job it is to map out and delineate our discipline. Beyond that, is the academy itself, whose traditions, standards, and requirements we have imbibed, deploying them rigorously in our own practice. And if we care to look still further, beyond the halls of academe, we may even see the society whose various institutions—from the logico-linguistic to the socio-political—make our investigations possible and before which our contributions will no doubt be held to account.
Cardiovascular diseases (CVDs), i.e., ischemic heart disease and stroke, remain the leading cause of death in the past decades around the world, especially in the developed countries [1]. CVDs can start from risk factors that may cause local vascular lesion and end up with systematic complications, which lead to organ failure and death. Thus, understanding the biochemistry of events involved in the whole process of CVD progression is crucial to prevent and treat the disease.
\nEpidemiological data show that various factors are associated with the increase of cardiovascular morbidity and mortality, including hypertension, smoking, hypercholesterolemia, diabetes mellitus, obesity, stress, low fruit and vegetable dietary, lack of regular exercise, and abnormal sleep [2]. Current therapeutic strategies mainly focus on reducing patients’ blood pressure, restoring redox balance, controlling cholesterol, and implementing physical activity programs [3]. In this chapter, we explore the physiological and pathological events in the cardiovascular system from the molecular biology’s perspectives. Molecular mechanism of muscle contraction and relaxation in the cardiovascular system will be discussed first. Then, we will delve into the biological effects of Nobel Prize molecule nitric oxide (NO), the most important vasodilator in the body. In addition, due to the inspiring clinical outcomes of using isosorbide dinitrate (an NO stimulus) and hydralazine (an antioxidant) to treat patients with symptomatic congestive heart failure [4], we will also discuss how nitroso-redox balance mediates cardiovascular functions.
\nSkeletal, cardiac, and smooth muscles have different structures and regulatory mechanisms, but they share the same molecular mechanism of contraction and relaxation, i.e., the relative sliding between myofilaments [5]. To understand how the heart beats and how blood vessels regulate their tones, it is important to look into the subcellular structure of these tissues (Figure 1).
\nMuscle contraction illustrated on different structural levels in the cardiovascular system: tissue level—heart (a) and blood vessel (b); cell level—cardiomyocyte (c) and vascular smooth muscle cells (SMCs) (d); subcellular level—filament sliding in cardiomyocyte (e) and SMCs (f); and molecular level—thin and thick filament for cross-bridge cycling (g).
Heartbeat relies on myofibrils, a fiber bundle structure that abounds in cardiomyocyte (Figure 1c). When a number of myofibrils are highly aligned, sarcomere, a repeat unit in the myofibril, can be observed under the microscope. Sarcomere is the basic unit for motion. Two most important proteins in the sarcomere are: myosin and actin. A myosin contains the N-terminal globular head domain, the short neck domain, and the long C-terminal coiled-coil tail domain. The globular head works as a specialized adenosine triphosphatase (ATPase), responsible for adenosine triphosphate (ATP) binding, actin binding, and generating force from ATP hydrolysis. The neck domain transduces force generated by the head. And the fibrous tails are bundled together to form the thick filament (Figure 1g). Actin, together with troponin and tropomyosin, forms the thin filament. When the two fibers slide toward each other, the overlapped region increases, which is the mechanism of muscle contraction. Similarly, when the fibers slide away from each other, muscles relax (Figure 1e).
\nThe filament sliding depends on cross-bridge cycling [6]. A cross-bridge refers to the two globular heads of myosin, which take turns to bind, pull, and detach from the actin fiber to achieve relative movement between the filaments. An analogy is that a person alternately uses two hands to pull a rope. One alteration of the hand is considered to be one cycle. There are four basic states [7] (sometimes detailed to six states [8]) in cross-bridge cycling. Each state corresponds to one behavior of ATP and one response of myosin. State 1: activation of myosin head, when ATP binds to myosin, it is hydrolyzed to ADP and Pi (inorganic phosphate); myosin becomes the “cocked position.” State 2: cross-bridge formation, the activated myosin binds to actin; Pi is released to stabilize the binding. State 3: power stroke, ADP is released; myosin generates force to pull actin filament. State 4: detachment of cross bridge, another ATP binds to myosin; myosin disengages from actin; then State 1 is repeated. The continuous cross-bridge cycling allows myosin to pull actin to its tail side, resulting in filament sliding and muscle contraction. At the resting state, actin’s myosin-binding site is blocked by troponin and tropomyosin [9] (Figure 1g). A switch mechanism is needed to expose and mask the myosin-binding site to regulate muscle contraction.
\nIntracellular Ca2+ works as a secondary messenger that quickly bonds to troponin, causing a quick conformational change of troponin and tropomyosin [10]. Thus, the myosin-binding site on actin filaments is exposed, and cross-bridge cycling proceeds. Intracellular Ca2+ concentration, or [Ca2+]i, can return to a very low level to cease the contraction and cause relaxation by different mechanisms, such as extruding Ca2+ out of cells or storing cytosolic Ca2+ into sarcoplasmic reticulum (SR) which functions as the Ca2+ reservoir in the cardiomyocyte. Similar mechanisms exist in the vascular tissue. SMC layer lies in between the endothelium layer and adventitia. There is no organized contractile protein fibril or sarcomere structure in SMCs [11]. Instead, the contractile fibrous proteins along with other intermediate filaments form bundles that are immobilized by anchoring proteins onto cell cytoskeletons. These filaments distribute all over the cytoplasm and connect each other through anchoring proteins (dense bodies) to form a three-dimensional network (Figure 1d and f). Unlike in cardiac muscles, actin filament in smooth muscles is associated with caldesmon, tropomyosin, and calmodulin (CaM) [12]. CaM is an important Ca2+ sensing protein, which binds and mediates many enzymes’ activities upon Ca2+ signaling. Caldesmon binds to actin, which inhibits the activity and motility of actin-myosin ATPase, and this binding is greatly strengthened by tropomyosin [13]. Ca2+ binds and activates CaM to uncouple the interaction between caldesmon and actin. Thus, actin’s myosin-binding sites are exposed to myosins. Different from skeleton or cardiac tissues, the contraction in smooth muscles also depends on phosphorylation level of myosin light chain, which is adjusted by the enzyme activity of CaM-dependent myosin light chain kinase (CaM-dependent MLCK) and myosin light chain phosphatase (MLCP) [14]. MLCK adds the phosphoryl group to the myosin light chain, while MLCP removes it. Thus, increase of [Ca2+]i also facilitates muscle contraction through enhancing myosin phosphorylation [15].
\nCardiovascular contractility is crucial for blood pressure homeostasis, thermal exchanging, mass transfer, immune responses, and organ functions [16]. Impaired coronary artery contractility (caused by the block of blood vessel or poor dilation) incurs ischemia heart disease [17]. To maintain cardiomyocyte viability and vascular tones, enhancing vasodilation and restricting oxidative stress are critical.
\nThe biochemical history of NO dates back to 1860s when the therapeutic use of nitroglycerin became a prevalent treatment for angina and hypertension [18]. Consequent studies showed that relaxation of blood vessels depended on a molecule present in an intact endothelium lining, which was named as endothelium-derived relaxing factor (EDRF). In 1977, it was demonstrated that NO was the chemical released from nitroglycerin metabolism [19], and EDRF was later identified to be NO [20]. Since then, NO’s cardiovascular effect and medical applications have drawn great research interest.
\nNO biosynthesis primarily relies on the enzyme nitric oxide synthase (NOS). The reaction uses substrate L-arginine and oxygen to generate L-citrulline and NO in the presence of the cofactors, including Ca2+/CaM, reduced nicotinamide-adenine-dinucleotide phosphate (NADPH), flavin adenine dinucleotide (FAD), flavin mononucleotide (FMN), and tetrahydrobiopterin (BH4) [21].
\nNO is an excellent messenger molecule in the human body because it is a highly reactive free radical and a highly diffusible molecule [22]. NO is able to react with various species, such as oxygen, superoxide, lipid oxygen radicals, thiols, and metals [23], which is the biochemical base for NO signaling. The ultimate biological effect of NO depends on its concentration, duration of release, and physiological environment [24]. NO concentration varies from subnanomolar (cell survival signaling) to micromolar (cytotoxic effect and apoptotic signaling) in the body [25]. Its small size and lipophilic characteristic allows it to move rapidly across cell membranes. However, NO’s effective distance is limited by its extremely high reactivity with species like oxygen, superoxide, and hemoglobin [23]. NO concentration decreases rapidly around the NO source, which makes NO’s effect extremely localized (within hundreds μm).
\nThree different isoforms of NOS have been identified in the human body. Endothelial NOS (eNOS) is mainly found in vascular endothelial cells (ECs) and is involved in regulating vascular tone and blood clotting. It is also present in cardiomyocytes where it mediates contractility of cardiac muscle [26]. Inducible NOS (iNOS) predominates in immune cells to produce large amounts of NO that aids in the host defense mechanisms. Examples include NO produced by macrophages and neutrophils when induced by lipopolysaccharide, interferon-γ, and tumor necrosis factor alpha [24]. Neuronal NOS (nNOS) is mainly identified in nerve cells where NO assists in nerve transmission. It has also been isolated from cardiomyocytes where NO regulates excitation contraction coupling of cardiac muscles [27]. NO generation by iNOS is mainly regulated at the transcription level, while eNOS and nNOS are constitutive NOS isoforms whose activities are Ca2+- and CaM dependent. Shear forces during blood flow stimulate the opening of Ca2+ channels on ECs to increase [Ca2+]i, resulting in eNOS activation [28]. NO biosynthesis can be inhibited by various chemicals that selectively bind to NOS with high affinities, such as NG-monomethyl L-arginine (L-NMMA) and asymmetric dimethyl L-arginine [9].
\nS-nitrosothiols and nitrite are alternative endogenous sources of NO. S-nitrosothiols decompose to release NO under physiological pH and are formed through the reaction of NO and thiol [29]. Nitrite can be reduced to NO in the body through pathways involving reducing agents and proteins, such as ascorbic acid, thiols, hemoglobin, and myoglobin [30]. These backup NO generation pathways are emphasized during hypoxia and acidosis when oxygen-dependent NOS-mediated NO production is limited.
\nEndogenous NO has various physiological effects in the body including inhibiting platelet aggregation, regulating SMC proliferation, modulating immune response, participating in neuron signal transmission, and inducing vasodilation [25]. In the vascular system, NO is primarily generated by eNOS in ECs (Figure 2). Shear stresses induced by blood flow and chemical stimuli, known as agonists including acetylcholine, bradykinin, adenosine triphosphate, estrogen, and vascular growth factors, are able to activate eNOS [28, 31]. NO is a highly dynamic molecule that diffuses fast in both aqueous and lipid environment (with diffusion coefficient
Biosynthesis of NO by eNOS and the biological effects of NO in the vascular system.
Cyclic-GMP binds and activates cGMP-dependent protein kinase (protein kinase G, or PKG), which can phosphorylate the downstream targets to trigger vasodilation (Figure 2). This signaling pathway can be terminated by removing cGMP through converting cGMP to GMP by various phosphodiesterases (PDE) [35]. There are many types of PDE in the human body, and they play critical roles in regulating cardiovascular function, adrenal steroidogenesis, and phototransduction [35].
\nInterestingly, protein kinase A (PKA, cAMP-dependent protein kinase) and PKG share very similar nucleotide binding domains. Many studies have shown that cGMP can activate PKA downstream pathways and cAMP can also cross-activate PKG [36]. This cross regulation between cGMP and cAMP pathways sometimes complicates NO signaling, which will be shown later.
\nDue to NO’s vasodilation effect, NO releasing drugs such as organic nitrate, and nitro- and nitroso compounds have been used for treating angina pectoris, congestive heart failure damage from ischemia–reperfusion, and pulmonary hypertension [37, 38]. Potent drugs also include chemicals that target members involved in NO-cGMP pathway. For example, Sildenafil, known as Viagra, is a PDE5 inhibitor. It prohibits cGMP from being hydrolyzed by PDE5 and extends the activation time of vasodilation to widen the blood vessel and increases blood flow into the penis to treat erectile dysfunction [39].
\nTo fully understand how NO causes vasodilation, it is necessary to perceive the relationship between NO-cGMP pathway and [Ca2+]i. At rest, extracellular Ca2+ concentration is high (1–2 mM), while the cytosolic Ca2+ is over 1000 times lower (>1 μM) [40, 41]. In the endoplasmic reticulum (ER, or SR in cardiac muscles), Ca2+ concentration is also high (about 400 μM) [40]. NO modulates [Ca2+]i by controlling Ca2+ exchange mechanisms on both cell and SR membranes.
\nVoltage-gated Ca2+ channels regulate [Ca2+]i through sensing electrical signals to allow Ca2+ entering the cell. High voltage-activated L-type channels are broadly found in the cardiovascular system. L-type Ca2+ channel inhibitors, such as dihydropyridines, phenylalkylamines, and benzothiazepines, are a major class of drugs for treating CVDs [42]. The opening probability of L-type Ca2+ channel can be lowered by PKG indirectly [43]. PKG phosphorylates the K+ channel and increases its opening probability to hyperpolarize the cell membrane [44, 45]. The hyperpolarized cell membrane can no longer send electrical signals to activate the Ca2+ channel, and therefore, Ca2+ influx is inhibited. Besides PKG, high NO level also directly activates K+ channels to achieve aorta relaxation in a cGMP-independent fashion [46].
\nCa2+ pumping ATPase located on the cell membrane also extrudes Ca2+ from the cytosol. It binds Ca2+ with a high affinity and forces Ca2+ out of the cell even when [Ca2+]i is very low to maintain the low [Ca2+]i level at rest [47]. PKG can stimulate the Ca2+ pump, initiating the expulsion of cytosolic Ca2+ [48].
\nUnlike Ca2+ pump, Na+/Ca2+ exchanger is more effective in quickly removing cytosolic Ca2+, but its Ca2+ binding affinity is low [47]. This mechanism is crucial for preventing cells from the cytotoxicity of an acute high Ca2+ concentration. The driving force for Na+/Ca2+ exchanger is the stored sodium electrochemical gradient created by Na+/K+ channels. PKG can activate the Na+/K+ channel to cause more Na+ accumulated to indirectly facilitate Ca2+ removal [49, 50].
\nCa2+ pump ATPase also resides on the ER responsible for the uptake of cytosolic Ca2+ into the ER. NO pathway regulates ER Ca2+ pumping through phosphorylation of phospholamban by PKG [51]. Mainly identified in cardiac tissues, phospholamban is an inhibitor of SR Ca2+ pump. Phospholamban is normally phosphorylated by PKA, which diminishes its inhibitory effect to Ca2+ pump [52]. Interestingly, in neonatal cardiomyocytes and vascular SMCs, NO pathway also demonstrated relaxation effect through differentially phosphorylating phospholamban [53, 54].
\nInosital 1,4,5-trisphosphate (IP3) is a critical messenger molecule that can induce Ca2+ release from the ER reservoir. IP3 receptor resides on the ER and works as a chemical-activated Ca2+ channel. NO-cGMP pathway can reduce IP3 generation [55], and PKG can phosphorylate and inactivate IP3 receptor in vascular SMCs to inhibit ER Ca2+ release [35, 56].
\nIndependent from NO-Ca2+ pathway, in SMCs NO also increases MLCP activity and limits MLCK activity, resulting in a dephosphorylation shift of myosin light chain phosphorylation balance [15]. Thus, myosin cross-bridge cycling is inhibited, causing smooth muscle relaxation.
\nAnatomic alterations in the cardiovascular structure directly deteriorate cardiovascular functions. NO is a multifunctional regulator for homeostasis in the cardiovascular system. An intact endothelial layer is the hub for NO generation. Pathological changes in NO generation can trigger various local flaws that may progress to be systematic cardiovascular issues with time.
\nDeviant NO level causes change of endothelial permeability, a key characteristic for mass transfer and extravasation. Interestingly, increase, decrease, and no change of vascular permeability due to the presence of NO have been reported. Using high concentration (millimolar level) of exogenous NO donor spermine NONOate decreased endothelial permeability in the
Vascular permeability is mainly determined by tightness of cell-cell junctions [60]. Tight junctions (TJs) and adherens junctions (AJs) are the most abundant interendothelial junctions. And both junctions are closely related to actin cytoskeleton dynamics [61] (Figure 2). TJs are composed of series of transmembrane proteins that anchor to the actin cytoskeleton to hold cells together. They seal the cells to maintain cell polarity and prevent the molecules from traveling through the space between cells. AJs consist of clusters of transmembrane protein cadherin, which is connected to actin cytoskeleton on its cytoplasmic side and binds strongly with cadherins residing on the neighboring cell membrane. These junctions are important for transmitting mechanical force between cells and reinforcing tissues. Since both junctions directly connect cytoskeletons, the cytoskeleton’s behavior will influence cell-cell junctions and thus control vascular permeability. When actin and myosin filaments undergo relative sliding to cause cell contraction, the cytoskeleton-associated membrane proteins will be pulled into the cells, and cell-cell junctions are disrupted. NO mediates cell contraction by adjusting [Ca2+]i. Therefore, deviated NO level may cause the change of cell-cell junctions [60].
\nAnother important downstream molecule of NO is vascular endothelial growth factor (VEGF) which has been extensively studied in cancer research due to its angiogenic effect. VEGF was initially considered as a vascular permeability factor, because it caused the formation of leaky capillaries [62], which is an important characteristic in tumor and retinopathy. Low NO level induces VEGF synthesis under normoxia through the transcription factor hypoxia-inducible factor 1 (HIF-1) [25, 63]. VEGF activates Src kinases, which further phosphorylate cadherin and elicit its internalization [64]. Once cells lose cadherin interactions, gaps between cells form and endothelial permeability is increased.
\nOne distinctive characteristic of vascular SMCs is the phenotypic plasticity. Two most important phenotypes are contractile and synthetic. Contractile SMCs guarantee the good performance of muscle contraction/relaxation, while synthetic SMCs are highly proliferative and migratory, crucial for vascular remodeling during pregnancy and injury healing. Dysregulation of the phenotype transition causes neointima formation [65]. NO plays important roles in suppressing SMCs’ contractile to synthetic transition.
\nNO donors and 8-Br-cGMP showed similar effect in inhibiting SMC migration and proliferation, indicating NO’s inhibitory effect might be through the cGMP-dependent pathway [66]. SMCs overgrow when stimulated by serum and epidermal growth factor (EGF). Many studies were based on these models, though divergent results were reported. EGF induces SMC proliferation through mitogen-activated protein kinases (MAPK) pathway, also called extracellular signal-regulated kinases (ERK) pathway. Ras (a small GTPase) and Raf (kinase of MAPK kinase, or MAPKKK) are the critical upstream protein kinases in this pathway. NO blocks MAPK pathway by prohibiting Raf from being activated by Ras-GTP in rat aortic SMCs. It is believed that PKG phosphorylates Raf, resulting in the conformation change. Thus, Ras-GTP cannot recognize Raf, causing the block of MAPK pathway and the accumulation of Ras-GTP [67]. Meanwhile, elevation of cGMP induced by IL-1β is correlated with the activation of PKA, and it can be prevented by blocking NO and cGMP pathways. Interestingly, this effect is cAMP independent, but PKA inhibitor, not PKG inhibitor, can prevent the inhibition of the proliferation, indicating that cGMP-PKA cross talk plays important roles in suppressing rat aortic SMCs’ proliferation [68].
\nNO-cGMP pathway may inhibit SMC growth by impairing cytoskeleton reorganization. Vasodilator-stimulated phosphoprotein (VASP) is characterized as a substrate of both PKG and PKA [69]. It targets focal adhesions and is involved in actin filament formation. Cell morphology change during proliferation relies on VASP, and its activation relies on the phosphorylation of Ser157 primarily mediated by PKA [70]. However, PKG can phosphorylate Ser239 and Thr278 to impair VASP’s activity and inhibit actin cytoskeleton reorganization [70, 71].
\nNO also directly mediates proteins associated with cell cycle and cell metabolism by cGMP-independent mechanisms. Cyclin A and cyclin-dependent kinase 2 expression levels can be blunted by exogenous NO donor DETA NONOate in an
Thrombus formation is critical for hemostasis during injury. However, thrombus in blood vessels can cause stroke and heart attack. Stable thrombus reduces lumen size and stiffens blood vessels. Unstable thrombus may rupture with blood flow and block the vessel. Activation of platelet is a critical step for thrombus formation, which involves exocytosis processes to expose P-selectin on the platelet surface and activate glycoprotein IIb/IIIa. Both processes depend on the elevation of [Ca2+]i controlled by IP3 pathway. NO suppresses platelet activation through NO-cGMP pathway [74]. Although, the inhibition pathway has not been fully characterized, evidences have shown that cGMP-PKG blocks agonist-induced IP3 formation in platelet [75], and PKG can phosphorylate IP3 receptor to inhibit Ca2+ release from the ER [35].
\nWhen the endothelium loses its integrity, there will be a local shortage of thromboregulators such as NO, prostacyclin, and ectonucleotidase CD 39, resulting in thrombogenesis [76]. Collagen and tissue factors also trigger the coagulation reactions [76]. The use of blood contact implant is another common source of thrombus. Note that, all materials are thrombogenic to some degrees. To enhance implant biocompatibility, an efficient method is to use NO releasing polymers to fabricate or surface coat the blood contacting devices (such as vascular graft/stent, intravascular catheter, and sensor implants). Common strategies include: physically incorporating NO releasing chemicals into polymer matrices, chemically linking NO releasing agent to polymer backbones, and developing materials that can trigger NO generation using endogenous NO donors circulating in the blood. By using the first two strategies, successful trials have been reported to achieve long-term (over few weeks to months) NO releasing and antithrombotic applications [77, 78, 79]. Good NO donors include N-diazeniumdiolate and S-nitrosothiols. Both hydrophilic and hydrophobic polymers that are commonly used in medical device fabrication have been successfully modified for NO release including poly(vinyl chloride), polymethacrylates, various hydrogels, polyethylene terephthalate, polyurethane, and silicone rubbers [77]. The third strategy directly uses endogenous NO donors as the NO reservoir to catalyze NO generation from S-nitrosoglutathione or nitrite in the body. Currently, its main challenge is to adjust the NO releasing rate to be more biologically relevant.
\nOxidative stress is always associated with ischemia reperfusion injury, dilated cardiomyopathy, and heart failure [80]. It is crucial to restore redox balance in the cardiovascular system when treating these diseases. Redox balance is governed by changes in the oxidative state in tissues, where addition and loss of electrons result in reduction and oxidation of molecules, respectively [80]. Oxygen can accept an electron to become reactive oxygen species (ROS). ROS are highly reactive chemical species that contain oxygen atoms, mostly free radicals with one or more unpaired electrons [81]. NO is a free radical signaling molecule. Under pathological conditions, it reacts with superoxide to generate reactive nitrogen species (RNS) that have detrimental consequences to cells. Herein, we highlight the causes of redox imbalance, their functions in the cardiovascular system, and the roles they play in the progression of CVDs.
\nThe electron transport chain (ETC) located in the inner membrane of mitochondria is crucial for energy and ROS generation (Figure 3). Normally, the final electron acceptor oxygen is reduced to water. However, in pathological conditions, electrons uncouple from the chain and react with oxygen without passing
The major sources of ROS and RNS in the cardiovascular system.
Superoxide is an anion-free radical that can produce other ROS including H2O2, hydroxyl radicals (.OH), and hypochlorous acid (HClO) [80, 82, 90, 91]. The spontaneous transfer of an electron to superoxide at low pH or by an enzyme reaction (superoxide dismutase, SOD) produces H2O2 [82, 91]. Low levels of H2O2 (1–10 nM) induce more antioxidant molecules that protect the cells, and high levels (>100 nM) are likely to generate more prooxidants [91]. For example, high level of H2O2 was generated in neutrophils for antimicrobial effects [92, 93]. Hydroxyl radical can be formed from the reaction between H2O2 and superoxide (Haber Weiss reaction) or the breakdown of H2O2 by metal ions, Fe2+ or Cu2+ (Fenton reaction) [94]. Hydroxyl radical is highly reactive. It alters DNA structure by attacking purine and pyrimidine bases, leading to mutations and cell damages [95]. In the pathological myocardial tissue, it is associated with decreased contractile function, increased membrane phospholipid peroxidation, and heart failure [96, 97]. HClO is mainly produced by leukocytes when H2O2 reacts with chloride anions. It facilitates the removal of foreign particles and is also implicated in the progression of atherosclerosis and ischemic reperfusion injury [81].
\nNO acts in a diffusion- and concentration-dependent manner. Low concentrations of NO (nanomolar range) have a protective role, while high NO levels (micromolar range) can be detrimental [98]. The majority of NO’s biological effect is attributed to sGC/cGMP pathway [21]. Additionally, NO acts as a signaling mediator through S-nitrosylation. NO can inhibit cardiac hypertrophy through nitrosylation of histone deacetylase 2 (HDAC2) released from chromatin [99]. HDAC2 regulates anti-hypertrophic genes. In ischemic preconditioning (the body’s defense mechanism against myocardial necrosis), the S-nitrosylation of mitochondrial proteins protects the mitochondria from oxidative stress [100]. S-nitrosylation initiates excitation contraction coupling by increasing Ca2+ uptake, and the contraction may be sustained through releasing of Ca2+ via SR membrane ryanodine receptors (RyRs) [26]. Quantitatively, when three thiols per subunit of RyR channels are nitrosylated, the process is reversible. However, if six or more thiols per subunit are nitrosylated, irreversible Ca2+ ion release occurs and can be detrimental to the cardiac muscle [26]. In addition, when too much NO is generated during inflammation or sepsis, NO may cause hypovolemia due to its excessive vasodilation effect [83]. Furthermore, upregulation of iNOS in ECs reduces the availability of BH4 to eNOS, intensifying eNOS uncoupling and superoxide generation [83]. Thus, the physiological role of NO can be attenuated by ROS, because NO is quickly consumed by superoxide before it initiates any cell response [101].
\nWhen NO collides with superoxide, peroxynitrite (ONOO−) is formed, causing nitrosative stress. The chemical reaction is very fast and deleterious [98]. ONOO− is a very strong oxidant. It reacts with proteins through tyrosine and tryptophan residues to form nitrotyrosine and nitrotryptophan, respectively [80, 98]. In diabetic mice, tyrosine nitration of the voltage-gated K+ channels in the vascular SMCs altered its dilation function, a possible mechanism of the progression of coronary artery disease [102]. Tyrosine nitration was also observed in cardiac myocytes desmin, myosin heavy chain, α-actin, and microtubules. These proteins play pivotal roles in maintaining cell morphology and cardiac contractility [98]. When free nitrotyrosine was incorporated into the carboxyl terminus of α-tubulin in microtubules, altered microtubule organization and redistribution of the motor cytoplasmic protein dynein were observed [103]. Protein activity can also be impaired by oxidation of thiols to disulfide bond by ONOO− [98]. In addition, ONOO− also reacts with lipids to yield nitrated lipids to promote atherosclerosis, and with nucleic acids via guanine and the sugar phosphate backbone to damage DNA [98].
\nOn the other hand, low concentrations of ONOO− (10–200 uM) is associated with tyrosine kinase-dependent signaling. ONOO− has been shown to activate tyrosine phosphorylation and trigger glycolysis [98]. Another example involves MAPK pathway, where ONOO− activates Raf-1 kinase. The MAPK pathway is closely associated with anti-apoptosis and cardiac hypertrophy in the cultured cardiomyocyte model [104].
\nAtherosclerosis is characterized by the formation of plaques that reduce the lumen of arteries and consequently interfere with blood flow and tissue perfusion. The plaque consists of the lipid core and fibrous cap. In patients with hypercholesterolemia, ROS and RNS oxidize low-density lipoprotein (LDL) [105]. Oxidized LDL (Ox LDL) initiates a cascade of events that alters the endothelial permeability and leads to insudation of the lipoprotein in the arterial wall. Stimulated by atheroprone signals, ECs express selectins and vascular cell adhesion molecule (VCAM-1) to attract circulating blood monocytes. Monocytes penetrate the endothelial layer; i.e., diapedesis occurs, and become macrophages [106]. Macrophages target Ox LDL for phagocytosis and become foam cells, the accumulation of which causes the formation of fatty streaks. The foam cells initiate the production of transforming growth factor beta (TGF-β), platelet-derived growth factor (PDGF), and fibroblast growth factor (FGF) in the vascular system [107, 108]. These growth factors promote the change of vascular SMCs from a contractile to a synthetic phenotype. SMCs migrate from the media layer to the intima, where they secrete a complex extracellular matrix to form a fibrous cap around the lipid core to stabilize the plaque [109]. The proliferation of SMCs leads to neointima hyperplasia. Thus, the vessel becomes narrowed and the blood flow profile alters, further aggravating endothelial dysfunction (Figure 4).
\nNO’s role in the initiation and progression of atherosclerosis.
In fact, disturbed blood flow at arches, branches, or bifurcations is always associated with the early appearance and fast development of atherosclerotic lesions. Blood flow influences ECs’ gene expression through “shear-stress response elements” in the promoters of atherosclerosis relevant genes and “mechanotransducers” that can sense the force and transduce mechanical signal into biochemical events within the cell. Overall, in steady laminar flow, ECs express more antithrombotic, anti-inflammatory, and antioxidant proteins, such as eNOS, cyclooxygenase-2 (COX-2), and manganese-dependent superoxide dismutase (SOD) [110], while in turbulent flow, ECs show atheroprone phenotypes, which activate NF-κB pathways to promote the expression of cytokines and cell adhesion molecules [107].
\nTwo highly differentially expressed transcription factors, zinc finger transcription factor Kruppel-like factor 2 (KLF2) and nuclear factor erythroid-2-related factor-2 (Nrf2), were identified by comparing endothelial gene expressions under different hemodynamic patterns [111]. KLF-2 maintains endothelial homeostasis at least in part by inhibiting local inflammation and restoring NO levels. Overexpression of KLF-2 blocks IL-1β-induced inflammation through inhibiting VCAM-1 and E-selectin expression to disturb the adhesion of immune cells [112]. In addition, it upregulates eNOS expression to improve vascular tones. Nrf2 is responsible for regulating redox-related genes (heme oxygenase 1, ferritin heavy chain, NADPH dehydrogenase quinone 1, and thioredoxin reductase) to maintain vascular redox balance in laminar flow [111]. Remarkably, it has been shown that KLF2 and Nrf2 work synergistically to integrate atheroprotective signals and active antioxidant responses, which may be a promising therapeutic strategy for CVDs.
\nTo counteract the effect of excessive ROS and control CVD symptoms, introducing antioxidative mechanisms is an effective method (Figure 5). Increasing enzymes that can eliminate ROS is a commonly used strategy. For example, superoxide can be eliminated by dismutation of two superoxide molecules by SOD to O2 and H2O2 [113]. H2O2 can undergo decomposition under the regulation of catalase and peroxiredoxin to oxygen and water [80, 114]. The thiol group in peroxiredoxins consumes H2O2 to form sulfenic acid, then subsequently disulfide bond [115]. Glutathione (GSH) peroxidase 1 uses the similar mechanism to inactivate H2O2, superoxide, and ONOO− in the presence of the tripeptide compound GSH. A prospective cohort study showed that reduced levels of GSH peroxidase 1 were associated with increased mortality in coronary disease patients [116].
\nMaintenance of redox balance in the cardiovascular system.
Another effective antioxidative method is to protect redox-sensitive molecules from being oxidized. In the body, the thiol group on GSH can form reversible mixed disulfide bonds with cellular proteins under oxidative stress conditions. These disulfide bonds can be broken by the enzyme glutaredoxin when the surrounding cell environment reverts back to its normal state [80, 117]. The addition of scavengers to directly remove ROS/RNS can also restore the nitroso-redox balance. An example is the elimination of superoxide by ascorbic acid (vitamin C) [113]. By limiting superoxide, other reactive species can also be repressed, such as •OH and ONOO−. This may explain the success of the clinical trial of combining nitrate drug isosorbide dinitrate with hydralazine, a NADPH oxidase inhibitor, where heart failure was reduced by 45% [118]. By inhibiting superoxide generation from NADPH oxidase, ONOO− level may be reduced and NO function preserved.
\nThe high concentrations of NO can be controlled through scavenging NO via oxyhemoglobin in red blood cells and myoglobin in the skeletal and heart muscle. These two proteins react with NO to form nitrate, which is considered as the primary method for inactivating NO in the cardiovascular system [119]. Hemoglobin and myoglobin can also scavenge ONOO− by their metal centers, generating nitrate from the reactions [120].
\nWe briefly reviewed the molecular mechanisms of muscle contraction and relaxation in the cardiovascular system and highlighted the importance of physiological and pathological effects of NO and oxidative stress. NO and ROS both determine the structural integrity and functionality of the cardiovascular system. The cardiovascular system not only nourishes cells, but also provides paths for immune response and systematic signaling. Drugs are transported by this system to the correct site for metabolic reactions. Tissue regeneration also relies on a healthy cardiovascular system. Therefore, to maintain, the homeostasis of the cardiovascular system is essential for overall health. Unfortunately, with aging, both cardiac function and cardiomyocyte number decline [121], and blood vessels undergo structural alterations [122]. Moreover, CVDs are also associated with other serious complications, such as diabetes, cancer, kidney failure, and inflammatory processes. Thus, multiple therapeutic strategies are needed to treat CVDs. According to 2011’s American Heart Association’s guidelines for preventing CVDs, therapeutic strategies include smoking cessation, blood pressure control, lipid management, physical activity programs, diabetes management, anticoagulation, dilation management, and depression prevention [3]. Besides traditional pharmaceutical management and surgeries, new perspectives to study, diagnose, and treat CVDs have also shown promising results, including development of biocompatible stents [123], stem cells therapies [124, 125], novel devices for mechanical thrombectomy [126], and inflammation management [127]. Although challenges still exist, the implementations of research findings from different disciplines in clinical trials will allow us to better understand and control CVDs in the future.
\nThe authors have declared that no conflict of interest exists.
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Fungal infectious illness prevalence and prognosis are determined by the exposure between fungi and host, host immunological state, fungal virulence, and early and accurate diagnosis and treatment. \r\nPatients with both congenital and acquired immunodeficiency are more likely to be infected with opportunistic mycosis. Fungal infectious disease outbreaks are common during the post- disaster rebuilding era, which is characterised by high population density, migration, and poor health and medical conditions.\r\nSystemic or local fungal infection is mainly associated with the fungi directly inhaled or inoculated in the environment during the disaster. The most common fungal infection pathways are human to human (anthropophilic), animal to human (zoophilic), and environment to human (soilophile). Diseases are common as a result of widespread exposure to pathogenic fungus dispersed into the environment. \r\nFungi that are both common and emerging are intertwined. In Southeast Asia, for example, Talaromyces marneffei is an important pathogenic thermally dimorphic fungus that causes systemic mycosis. Widespread fungal infections with complicated and variable clinical manifestations, such as Candida auris infection resistant to several antifungal medicines, Covid-19 associated with Trichoderma, and terbinafine resistant dermatophytosis in India, are among the most serious disorders. \r\nInappropriate local or systemic use of glucocorticoids, as well as their immunosuppressive effects, may lead to changes in fungal infection spectrum and clinical characteristics. Hematogenous candidiasis is a worrisome issue that affects people all over the world, particularly ICU patients. CARD9 deficiency and fungal infection have been major issues in recent years. Invasive aspergillosis is associated with a significant death rate. Special attention should be given to endemic fungal infections, identification of important clinical fungal infections advanced in yeasts, filamentous fungal infections, skin mycobiome and fungal genomes, and immunity to fungal infections.\r\nIn addition, endemic fungal diseases or uncommon fungal infections caused by Mucor irregularis, dermatophytosis, Malassezia, cryptococcosis, chromoblastomycosis, coccidiosis, blastomycosis, histoplasmosis, sporotrichosis, and other fungi, should be monitored. \r\nThis topic includes the research progress on the etiology and pathogenesis of fungal infections, new methods of isolation and identification, rapid detection, drug sensitivity testing, new antifungal drugs, schemes and case series reports. It will provide significant opportunities and support for scientists, clinical doctors, mycologists, antifungal drug researchers, public health practitioners, and epidemiologists from all over the world to share new research, ideas and solutions to promote the development and progress of medical mycology.",coverUrl:"https://cdn.intechopen.com/series_topics/covers/4.jpg",keywords:"Emerging Fungal Pathogens, Invasive Infections, Epidemiology, Cell Membrane, Fungal Virulence, Diagnosis, Treatment"},{id:"5",title:"Parasitic Infectious Diseases",scope:"Parasitic diseases have evolved alongside their human hosts. In many cases, these diseases have adapted so well that they have developed efficient resilience methods in the human host and can live in the host for years. Others, particularly some blood parasites, can cause very acute diseases and are responsible for millions of deaths yearly. Many parasitic diseases are classified as neglected tropical diseases because they have received minimal funding over recent years and, in many cases, are under-reported despite the critical role they play in morbidity and mortality among human and animal hosts. The current topic, Parasitic Infectious Diseases, in the Infectious Diseases Series aims to publish studies on the systematics, epidemiology, molecular biology, genomics, pathogenesis, genetics, and clinical significance of parasitic diseases from blood borne to intestinal parasites as well as zoonotic parasites. We hope to cover all aspects of parasitic diseases to provide current and relevant research data on these very important diseases. In the current atmosphere of the Coronavirus pandemic, communities around the world, particularly those in different underdeveloped areas, are faced with the growing challenges of the high burden of parasitic diseases. At the same time, they are faced with the Covid-19 pandemic leading to what some authors have called potential syndemics that might worsen the outcome of such infections. Therefore, it is important to conduct studies that examine parasitic infections in the context of the coronavirus pandemic for the benefit of all communities to help foster more informed decisions for the betterment of human and animal health.",coverUrl:"https://cdn.intechopen.com/series_topics/covers/5.jpg",keywords:"Blood Borne Parasites, Intestinal Parasites, Protozoa, Helminths, Arthropods, Water Born Parasites, Epidemiology, Molecular Biology, Systematics, Genomics, Proteomics, Ecology"},{id:"6",title:"Viral Infectious Diseases",scope:"The Viral Infectious Diseases Book Series aims to provide a comprehensive overview of recent research trends and discoveries in various viral infectious diseases emerging around the globe. The emergence of any viral disease is hard to anticipate, which often contributes to death. A viral disease can be defined as an infectious disease that has recently appeared within a population or exists in nature with the rapid expansion of incident or geographic range. This series will focus on various crucial factors related to emerging viral infectious diseases, including epidemiology, pathogenesis, host immune response, clinical manifestations, diagnosis, treatment, and clinical recommendations for managing viral infectious diseases, highlighting the recent issues with future directions for effective therapeutic strategies.",coverUrl:"https://cdn.intechopen.com/series_topics/covers/6.jpg",keywords:"Novel Viruses, Virus Transmission, Virus Evolution, Molecular Virology, Control and Prevention, Virus-host Interaction"}],annualVolumeBook:{},thematicCollection:[],selectedSeries:null,selectedSubseries:null},seriesLanding:{item:null},libraryRecommendation:{success:null,errors:{},institutions:[]},route:{name:"chapter.detail",path:"/chapters/64961",hash:"",query:{},params:{id:"64961"},fullPath:"/chapters/64961",meta:{},from:{name:null,path:"/",hash:"",query:{},params:{},fullPath:"/",meta:{}}}},function(){var e;(e=document.currentScript||document.scripts[document.scripts.length-1]).parentNode.removeChild(e)}()