Context of denture and overdenture complications – key factors
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Dentures and overdentures, the most frequently used treatment options for the complete edentulism, can have local and systemic complications. For their prevention, treatment and reduction of their negative impact, it is necessary to understand their etiological context and to know their particularities of manifestation. Considering the relatively high rate of some complications of denture and overdenture treatment, knowing them is essential for ensuring a treatment that corresponds to the medical standards of care and patients’ needs and expectations.
All medical treatments should be approached with a holistic perspective in mind, due to the fact that there are numerous factors which, through interacting each other, have an impact on the final medical outcome. Understanding the problem and its realistic possible approaches, but also considering its treatment limitations and performing an analysis that evaluates the medium and long-term prognosis ensures the highest premises for obtaining a good result.
The previous also applies to the treatment of edentulism using dentures or overdentures. Some of the key aspects that might help understand better the denture and overdenture complications, as they define the etiological context, are mentioned in Table 1.
Context | \n\t\t\tGeneral medical and social factors | \n\t\t\tMedical and social perception of edentulism Demographics of edentulism | \n\t\t
Denture and overdenture treatment factors | \n\t\t\tTreatment difficulty Treatment options overview Maintenance therapy Technical and biomechanical considerations Previous dental treatments | \n\t\t|
Edentulous patient factors | \n\t\t\tOral health status Systemic health status and medication use Age Health risk factors Patient need and preferences | \n\t\t
Context of denture and overdenture complications – key factors
Edentulism is defined as the loss of all permanent teeth. Tooth loss is an outcome of a complex interaction between disease entities (e.g., caries and periodontal disease) and non-disease entities (e.g. economy, oral healthcare system, access to dental services, dental awareness, cultural tradition, education) [1]. Continuing exposure to risk factors after onset of edentulism (e.g., poor oral hygiene, smoking, deficient dental treatment) can have an etiological role in the occurrence of complication.
Edentulism is a chronic, severe, irreversible medical condition and is described as the final marker of disease burden for oral health [2,3]. It is common for elderly people, but it is not regarded any more as an inevitable phenomenon that comes with age [4].
Edentulism has several deleterious consequences on oral health (e.g., residual ridge resorption, impaired masticatory function, trouble speaking), general health (deficient nutritional status, increased risk for certain systemic diseases), mental and social well-being (dissatisfaction with appearance, avoidance of social contacts) and on quality of life [1,2,4]. The previous have impact on prosthetic treatment to be performed.
Thus, the current perception on edentulism is as non-fatal sequelae of diseases and injuries, which still represents a tremendous global health care burden [5,6]. It can be considered a physical impairment, because important body parts have been lost, a disability, because it associates functional limitations or a handicap, as it sometimes limits or prevents normal life or work activities [1,2,7-9].
Considering the impact and demographics of the edentulism, the health care barriers that older people face, the Active Ageing approach of the World Health Organization (keeping older people socially engaged and productive), intensive measures and new regulations regarding caring for the elderly population are needed. Consequently, implementation of gerodontology, as a new dental specialty, may be appropriate [10].
According to the current reports and predictions, edentulism is and will continue to represent a common disease for the elderly people segment.
There is a tendency for reduction of the edentulism prevalence, through the reduction of tooth loss. Thus, in the United States in the period of 1999-2004, the prevalence of tooth retention in seniors (65 years and older) significantly increased from 17.9 teeth to 18.9 teeth and the prevalence of edentulism significantly decreased from approximately 34% to 27% [11]. This phenomenon can be justified through the progress made in the dental field, the emphasis on prevention measures, improved access to dental care services and mass education for approaching a healthy behavior [4]. But, despite these efforts, complete edentulism continues to have a high prevalence, aspect associated mainly to the aging population phenomenon through growth of the life expectancy and thus the number of elderly people and the number of edentulous patients [12,13].
Estimates show that edentulism is found in 2.3% of the world’s population regardless of age, respectively in 7-69% of adult populations internationally [5,14]. Considerably high disparities are noted between different countries, different regions, due to the important impact of the socio-economical and behavioral factors.
The prognoses show that edentulism is decreasing, but most probably will continue to be a condition with a significant prevalence, especially in elderly’s people, which is estimated to be a growing category in the global population [15]. Douglas estimates that in the United States the population with one or two edentulous jaws will increase from 34 million in 1991 to 38 million in 2020 [1,12]. Felton considers that most probably the necessity for complete denture therapy will not disappear over the next 4 or 5 decades, and the economic conditions may even lead to a growing need [1,6].
Edentulism is generally regarded as a clinical condition with a high degree of treatment difficulty, often being hard to achieve optimal functional parameters. The complexity of the edentulous condition derives from the extensive oral changes, both anatomical and functional, that sometimes require preprosthetic surgical intervention in order to optimize the biomechanical conditions, which are superimposed on general alterations (related to age, systemic disease, and psychosocial status). In order to support the differentiation of cases according to their treatment difficulty degree, the ACP (American College of Prosthodontists) has put together the Prosthodontic Diagnostic Index (PDI) Classification System for the complete edentulism [16]. Higher complexity of edentulism condition increase the risk of treatment complication (e.g., in cases with severe ridge resorption ill-fitting dentures are more frequently noticed), and complications can also contribute to increasing the degree of treatment difficulty (e.g., wearing unstable dentures accelerate the ridge resorption rate).
Complete denture used to be the only treatment option for the complete edentulism. Nowadays, this is still the most frequently used treatment option, but there can be seen a growing trend towards using implant prosthetic restorations fixed or removable. Each treatment option has the risk of specific complications, dependent on their manufacturing particularities and bio-mechanical features.
Dentures can have both local and systemic complications, such as gingival hyperplasia, denture stomatitis, loss of denture retention, fracture of the denture and functional impairment, mastication deficiencies having a negative impact on the nutritional status. Some patients cannot tolerate the dentures, aspect that can be connected to psychological factors, to patients’ needs and expectations, but also to age, oral conditions, denture deficiencies and doctor-patient relationship.
Root supported overdentures, with or without attachment systems, have the advantage of improved retention and stability, with a positive impact on the oral functions and the accommodation with the future dentures. Their possible complications include the ones of the conventional dentures and, additionally, some modifications of the supporting teeth or the attachment system used.
Prosthetic implant restorations, either fixed or a removable, are alternatives that provide an improved functional integration and better treatment outcome, but are more complex and require preprosthetic interventions, with additional biological, financial and time costs. Using these treatment options involves the risk for additional complications, with regards to the higher complexity of the treatment –e.g., treatment plan related, surgical complications, technical complications.
Maintenance is very important for the longevity of the treatment, having a positive impact in reducing the frequency and severity of its complications. Both type of procedures, those performed in the dental office, by the dentist and at home, by the patient, are relevant in this respect.
Periodical check-ups are essential, considering that there are some complications with a high prevalence rate both for dentures and implant overdentures (e.g., loss of denture stability due to progressive ridge resorption, denture adjustments and relinings, clip activations) [17]. Additionally, the edentulous patients are often elderly patients, and face access barriers to dental care services, in relation to aspects like lack of finances or transportation difficulties [18,19]. Due to this, it is recommended to keep in mind the possible complications and to take the appropriate preventive measures to limit them at the time the treatment is being planned and performed.
Informing and instructing the patient on how to take proper care of the oral care and prosthetic restorations are important aspects, since complications can be tightly related to this (e.g., the lack of appropriate cleaning of the denture, teeth or implants is associated with a higher risk for denture stomatitis, tooth or implant loss). Since we are frequently dealing with elderly people, who have less manual dexterity, it is recommended to choose simpler treatment option (e.g., if applicable, 2-implant overdentures are more appropriate than 4-implant overdenture [20].
According to the current level of knowledge, treatment with dentures or implant/root overdentures must consider the risk for developing complications in relation to the technical and biomechanical features (e.g., design, attachment components, materials).
There are different types of design for dentures and overdentures, with different possible complications. Thus, using narrow-diameter implants associates a higher risk of implant fracture. Considering the occlusal scheme, there is evidence that patients prefer dentures with lingualized occlusion [21]. Metal or non-metal (glass and polyethylene fibers) inserts are recommended for denture base reinforcement when there is a high risk of denture fracture or when there are more than 2 teeth or implants supporting the denture [22].
Material used for denture/overdenture fabrication associates the risk of developing complications in relation to their physico-chemical properties and their biocompatibility. For example, polymethylmethacrylate (PMMA), the material mostly used for manufacturing of dentures or overdentures, through its features (porosity, increased wettability, low mechanical strength, monomer release after curing) facilitates the occurrence of complications such as microbial or contact denture stomatitis, fracture of the dentures, artificial teeth discoloration and wear [23].
A key element in order to achieve a predictable outcome is the analysis of the previous dental and prosthetic treatments, by connecting patient’s subjective complaints with prosthetic restoration’s objective deficiencies. This gives important information that could be used for decision making in establishing the particularities of the future prosthetic treatment. For example, complete denture intolerance can be linked to personality traits, to objective patient’s features that enhance the occurrence of functional deficiencies, or to some objective faults of the dentures. Differentiating between these three situations is the basis for selecting the optimal treatment option, with the possibility to prevent the complications that occurred in the past.
The complete edentulism cannot be regarded simply as the loss of teeth. It is accompanied by massive, progressive changes of the oral structures and functional alterations, which associates a high degree of treatment difficulty and the occurrence of specific complications. Impact of edentulism on oral health is mainly manifested in 3 directions: modifier of normal physiology; risk factor for impaired mastication; determinant of oral health [2]. Amongst the sequelae of treatment with complete dentures, as the most commonly used treatment option, there can be mentioned residual ridge resorption, mucosal reactions, burning mouth syndrome, denture stomatitis [24].
Considering the severe changes of the oral status in edentulous patients, the increasing elderly population and the relatively frequent barriers to oral health care of older people (e.g., financial hardship, transportation difficulties), Petersen et al. makes a series of recommendations among which are the incorporation of age related oral health concerns into the promotion of general health, that could ease the development of oral health care for older people [25].
Between oral health and general health there are numerous interactions, that sometimes materializes as local or systemic complications.
The impact of complete edentulism on the general health status is manifested as an increased risk of conditions, such as nutritional deficiencies, inflammatory changes of the gastric mucosa, peptic or duodenal ulcers, obesity, noninsulin-dependent diabetes mellitus, hypertension, heart failure, ischemic heart disease, stroke, aortic valve sclerosis, chronic kidney disease, sleep-disordered breathing, including obstructive sleep apnea [2]. Additionally, functional limitations, mental and social well-being alterations that negatively impact the quality of life are more common in edentulous patients.
The impact of general health status and the medication used on the oral health of the edentulous patient is partially manifested through the occurrence of complications. Nutritional deficiencies increase the risk of occurrence of denture stomatitis, traumatic ulcer and burning mouth syndrome [25]. Patient’s personality and psychological well-being influences treatment satisfaction and tolerance [10]. Decreased manual dexterity has a negative impact on care and maintenance of dentures/overdentures, which leads to negative effects on oral and systemic health [14].
Patient’s age is an important aspect to consider when planning the prosthetic treatment, being linked to particularities of oral and general health status, to specific needs and expectation towards the prosthetic rehabilitation, to particular medical approaches in order to ensure a good long-term prognosis. Prosthetic treatment of the edentulous patient should take into account the current situation, but also the most probable evolution and, if present, the inherent complications (e.g., preventive measures to reduce alveolar ridge resorption are recommended).
Young-elderly edentulous patients generally have more favorable clinical conditions for prosthetic rehabilitation, a better general health status, a faster adaptation to removable prosthesis if chosen and the ability to perform most accurately the necessary the maintenance procedures. They have higher expectations regarding the esthetics and functionality of the prosthetic rehabilitation and don’t easily accept the removable treatment options.
Old-elderly edentulous patients generally register an increased treatment difficulty, as a consequence of numerous factors interacting. In previous ill-fitting complete denture wearers there is a severe ridge resorption [26,27]. The prevalence of co-morbidities is increased, such as physical or mental health problems that have a negative impact on oral health, systemic health, functioning and behavior. Most of the times the elderly people are not regular users of dental services since they overcome physical and psychological access treatment barriers (e.g., the cost of dental care services, transportation problems, doctor’s attitudes-lack of responsiveness to patient’s concerns, the lack of perceived need for care, fear), which are more significant for the functionally dependent elderly then for the independent elderly [28-30]. They have treatment expectations that target first the rehabilitation of the masticatory function, and second the esthetics. They usually prefer more simple medical procedures, that include limited surgical interventions and that demand easy maintenance procedures. The older completely edentulous patients show a more frequent rate of denture intolerance, probably due to less adaptability to new situations.
Demographic changes, namely population ageing and decreasing prevalence of tooth loss, have impact on the edentulous patient profile. There is an increasing of the age when edentulism occurs, aspects that associates an increased treatment difficulty. Considering the latter, additional measures are necessary to ensure adequate oral health for older edentulous patients e.g., access to and financing for dental services, an adequately trained workforce to provide dental care and appropriate education to edentulous individuals [30].
Health risk factors should be assessed since they can explain some of the case particularities and may have a negative impact on the treatment outcome. Among them, there can be mentioned behavioral risk factors (e.g. tobacco and alcohol consumption, obesity related to physical activity and diet), social risk factors (e.g., socio-economical status, social networks and social support, occupational factors, social inequalities), inadequate disease screening practices, exposure to increased stress [31]. Their role is proven both as a cause of complete edentulism and also as a factor that impacts the treatment outcome, being risk factors for some complications.
Health care decisions require integrating the patient’s individual preferences and values, according to the ethical principle of respecting the patient’s autonomy [32]. A good relation and communication between doctor and patient offers the best premises for reaching a consensus regarding the medical decision, with a positive effect on the treatment outcome.
Patient preferences are related to numerous variables, e.g., age, social status, personality type, education. Acknowledging them may be difficult, especially in elderly patients, sometimes in relation to objective reasons, as physical changes that affect the communication (e.g., loss of hearing or visual acuity). Additional efforts should be made in order to understand the patient’s health needs and preferences, since they can have important consequences, such as rejection of the prosthetic treatment or even avoiding addressing for medical treatment.
The classification of denture and overdenture complications can enhance practitioner’s understanding of them, with a positive effect on their management and prognosis.
Denture and overdenture complications can be classified considering their etiology, according to risk factor’s nature and mechanism of action, as described in table 2, or in regarded to some descriptive criteria, as presented in table 3.
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Classifications of denture and overdenture complications, considering their etiology
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Classifications of denture and overdenture complications, considering descriptive criteria
Some of the most common complications of the completely edentulous patient, treated by dentures or implant/root overdentures will be presented. Aspects related to their etiology, clinical features and management will be covered.
The residual ridge derives from the alveolar process after tooth loss. It registers the most significant changes and it supports the highest pressures during the worn of dentures or implant-retained overdentures. The ridge resorption is manifested as a continuous, cumulative and irreversible process, visible as the decrease of the quantity and quality of the bone [36].
The process of postextractive bone restructuring, after tooth loss, has variable rate and pattern, in relation to general physiological and pathological factors (age, menopause, systemic alterations), local factors (the edentulism and its cause, features of the jaws – volume, density). Also, the rate of bone resorption (the quantity of bone lost in a time period) varies in relation to the moment of tooth loss-it is maximum immediately after it in the first month, high in the first year after the tooth loss and decreases consequently. The pattern of bone resorption registers topographic differences – as for the maxilla and the mandible, for the anterior and posterior regions and in relation to anatomical features. The resorption is maximum at the top of the ridge and is lower at the base of the ridge, in the biostatical areas (maxillary tuberosity, retromolar pad), at the ligaments’ insertion site (frenum) and in the region of the hard palate. The ridge resorption occurs from the top to the basis, and is centripetal in the maxilla and centrifugal in the mandible. The pattern of ridge resorption varies according to the anatomical features and the size of the jaws, e.g., in class II skeletal patients, brachicephals, with mandibular micrognathism the resorption is more severe in the mandible, and in class III skeletal patients, dolicocephals, with mandibular macrognathism the resorption is more severe in the maxilla. Also, ridge resorption is more pronounced in women (probably linked to smaller jaws and lower bone density, related to postmenopause osteoporosis), in patients who lost their teeth due to periodontal disease and in those with high occlusal forces (natural teeth as antagonists, bruxism). Systemic conditions, particularly diabetes mellitus and other metabolic disorders, can accelerate the rhythm of ridge resorption.
The dentures accelerate the rate of ridge resorption, mainly through the pressure exercised by them on the support structures during oral functions. The severity of ridge resorption is connected to the parameters of functional and parafunctional forces of occlusion and to biomechanical aspects related to the prosthesis-the support and stability of the denture, the positioning of artificial teeth, type of occlusion, antagonists (teeth, implants, edentulous), and correctness of the registration of maxillomandibular relationship. The support surface for occlusal forces is reduced in edentulous patients, compared to the dentulous ones, and through progressive ridge resorption, both in high and width, consequently the support surface decreases even more. The magnitude of occlusal forces are generally lower in the edentulous patients, but there are variations related to age, sex, parafunctions as bruxism, stress level, food consistency preferences, and also the correctness of prosthetic rehabilitation. Increased duration of occlusion contacts, as a risk factor for ridge resorption, is related to bruxism, ill-fitting dentures, unstable occlusion and increased vertical dimension of occlusion. Compared to maxillary edentulism, mandibular edentulism has greater risk of registering more severe ridge resorption, due to the decreased denture support surface and related higher magnitude of pressure beared. Also, denture wearing associates the risk of specific complications that favor the occurrence of an accelerated rate of ridge resorption, such as inflammatory lesions of the oral mucosa (e.g., denture stomatitis). Due to these factors, it is considered that ridge resorption is in tight relation with the period of wearing the dentures, but is also influenced by the quality of the treatment.
Ridge resorption implies a decrease in bone volume, as ridges’ height (assessed as reduced, medium and severe resorption), ridge’s width (assessed as wide, medium or thin “knife edge ridge”) and ridge’s surface layout (normal or abnormal morphology, with exostosis). The characteristics of the alveolar ridge influence treatment conduct and have impact on its outcome, e.g., severe ridge resorption (Figure 1) is more frequently associated with denture instability and reduced denture tolerance, difficulties in mounting the artificial teeth and esthetic deficiencies.
Severe ridge resorption, in long-term denture wearers
Associated to ridge resorption particular aspects of the maxillomandibular relationship can be noticed, as lack of parallelism between the ridges direction and anterior or/and posterior inverse ridge relationship (Figure 2). According to their skeletal jaw relations and in relation with the different patterns of jaws resorption, class III skeletal patients have the tendency to register an inverse ridge relationship, and class II skeletal patients an apparently normal relationship.
Inverse ridge relationship, related to skeletal class III and the pattern of bone resorption (centripetal in the maxilla and centrifugal in the mandible)
Through resorption and replacement of the bone with fibrous tissue, a floating ridge, usually named “flabby ridge” is noticed. This aspect is most commonly observed in the edentulous anterior maxilla, being related to the excessive pressure of the mandibular anterior teeth (Combination Syndrome). Flabby ridge can also be seen in other places, like maxillary tuberosity or retromolar pad, being linked to instability of the denture or excessive occlusal trauma.
Severe mandibular ridge resorption is accompanied by reduction of the area of the fixed mucosa, difficulties in acknowledgement of the extension of the denture base (due to the sublingual gland herniation through the mylohyoid muscle and modifications of the muscle and ligaments’ insertion sites, which can get close to the ridge crest) and pain as a result of dental pressure in the mental foramen area and nerve exposure.
Denture wearing associates inherent ridge resorption, manifested as the occurrence of denture instability. Consequently, clinical procedures as relining or rebasing are required for readjustment of the dentures, in order to correspond to patient’s need and to prevent worsening of the edentulous condition.
In edentulous patients, considering the irreversible and progressive character of bone resorption, preventive interventions should be taken towards reduction of resorption rate and its complications. In this respect, addressing the risk factors and correct management of the supporting tissue should be a priority. In order to limit the bone resorption it is recommended to preserve the tooth roots, to use dental implants, to realize immediate prosthetic rehabilitation, especially in cases with tooth lost due to periodontal disease since this conduct favors a more reduced guided bone resorption. Correctness of dentures manufacturing is essential and it should rely on the principles of retention, stability and support, with proper maintenance and on time replacement. Implant overdentures can be used both as a preventive solution, in order to reduce the bone resorption, and as a curative solution, for solving the cases with severe ridge resorption where conventional dentures did not succeed or were not tolerated.
Severe ridge resorption associates decreased denture stability, which is associated with complications such as pain, lesions of the mucosa, reduced denture tolerance, that need to be addressed. The surgical preprosthetic interventions (bone augmentation, frenectomy, excision of hyperplasic lesions, as in figure 3) and non-surgical interventions (tissue conditioning, antifungal medication, improvement of the nutrition) are preparative treatments that aim achieving better conditions for prosthetic rehabilitation. Taking into account edentulous patient’s profile (aged, with systemic co-morbidities), stress related to the fear of surgical interventions and healing parameters (as time needed or remaining scar tissues), the non-surgical or less invasive surgical interventions are preferred. Soft lining materials are indicated since they facilitate the uniformly distribution of the functional stress and can reposition the abused tissues.
The prosthetic treatment of the edentulous patient can be performed using conventional or implant restorations, fixed or removable, with or without preprosthetic interventions, according to the clinical case’s particularities and patient’s needs. Treatment requirements include accurate physiological impression of the oral structures, correct registration of maxillomandibular relationship and teeth mounting and selection of appropriate occlusal scheme, in order to ensure dentures’ stability and esthetic and functional rehabilitation.
Preprosthetic surgical interventions for excision of hyperplastic lesions
Accurate establishment of the peripheral extension of the denture base, considering also the pressures supported by the denture-bearing area, is extremely important, being directly relate to denture’s retention, stability and tolerance. In this respect, the correct 2-phase impression technique (primary and custom tray impression) is essential. In edentulous patients with severe ridge resorption, additional adjunctive procedures may be required as tissue conditioning, supplementary functional impressions or usage of neutral zone impression technique. In displaceable or “flabby ridges”, the selective pressure impression technique (e.g., using a custom tray with a window opening over the mobile tissue) is more recommended, being at equal importance to other aspects as stable posterior occlusion. Thin mandibular “knife edge ridges”, that are accompanied by pain related to denture pressure, needs special treatment conduct, with usage of soft liners, a selective pressure impression technique, preprosthetic surgery (some disagree because ridge reduction implies loss of potential stabilizing zone) and dental implants.
Registration of maxillomandibular relationship is essential for the treatment success. It implies establishing the functional vertical dimension of occlusion, in accordance with minimum speaking space and the freeway space, and respecting the coincidence of maximal intercuspal position and centric relation. The most recommended occlusal schemes for removable prosthesis are the lingualized occlusion, for the bimaxillary complete edentulous patient, in skeletal class II patients or in severe mandibular ridge resorption or the linear occlusion, for mandibular overdentures, in patients with combination syndrome or skeletal class III pattern and severe maxillary ridge resorption.
Mandibular conventional dentures register frequently retention and stability deficiencies, mainly related to ridge resorption. These can be addressed through usage of implant prosthetic restorations, fixed or removable. There are multiple treatment options when considering usage of dental implants, as removable prosthesis (conventional or narrow dental implant overdenture, with different attachment systems as bars, ball, Locator) or fixed restorations (All an four, Fast & Fixed, conventional fixed implant restorations). Current perspective identifies 2 implant overdentures as the minimum standard for mandibular edentulism taking into account performance, patient satisfaction, cost and clinical time [37]. Selecting between them require acknowledgement of case futures and patient’s need and preferences. For example, fixed restorations have better treatment outcome, but have limited usage due to aspects like cost and higher complexity of the interventions required (e.g., sometimes surgical procedures as bone augmentation or sinus lift cannot be avoided).
Traumatic ulcers are small, painful mucosal lesions that most commonly develop in the first days after insertion of a new denture [38].
Denture related hyperplasia is an enlargement of the oral mucosa, appeared in relation to the denture base. There are two main types of denture related hyperplasia, namely denture-related fibrous hyperplasia (epulis fissuratum) and inflammatory papillary hyperplasia.
In inflammatory papillary hyperplasia the hard palatal mucosa has an erythematous aspect, with a pebbly or papillary surface [42]. According to its severity, we can see forms with limited localization or that cover the entire hard palatal mucosa. The previously described two types of denture related hyperplasia can be observed in figure 4.
Denture related hyperplasia
The treatment of inflammatory papillary hyperplasia requires removal of the denture at night, improvement of the oral hygiene and denture hygiene. Antifungal therapy, surgical excision of the hyperplastic tissues and renewal of the denture can be recommended in some cases [42].
Denture stomatitis is a chronic infectious inflammatory disease of the oral mucosa that is in direct contact with the base of the removable prosthesis, either conventional or implant-supported.
Acrylic dentures produce ecological changes that facilitate the accumulation of bacteria and yeasts and thus commensal organism may become pathogenic, denture stomatitis being considered an opportunistic infection [44]. A higher prevalence is noticed in cases with poor denture hygiene with denture plaque accumulation, continuous wear of the dentures (including at night) and in ill-fitting dentures. Other risk factors for denture stomatitis are related to the material characteristics, as their changes in time that favor plaque accumulation and microbial colonization (soft linings materials through their fast deterioration and difficulties of achieving proper hygiene; hard acrylic materials through their increased porosity that occurs in time) or as determining hypersensitivity reactions.
Host related risk factors for denture stomatitis include local factors (reduced salivary flow rate, low salivary pH, poor oral hygiene), general factors (physiological such as age, sex, nutritional status and associated medication) which act towards decreasing the resistance and defense mechanisms of the oral mucosa. The prevalence of denture stomatitis is higher among elderly denture users, women, smokers, alcohol consumers, vitamin A deficiency, diabetes and immune deficiency [44-47]. Changes in the salivary flow rate may be signs of a systemic disease, as in Sjögren or Mikulicz syndromes, or associated to medication use, as diuretics, antihypertensive, antipsychotic, anxiolytic, analgesic, anti-inflammatory, antihistaminic drugs. Also, incorrect antibiotic therapy, without fungal protection and broad spectrum antibiotics are seen as risk factors.
Denture stomatitis – clinical aspect
Treatment of denture stomatitis consists mainly in adopting strict methods for oral and denture hygiene, with removal of the denture overnight and soaking it in an antiseptic solution, such as chlorhexidine mouthwash. Considering the frequent Candida colonization, antifungal agents, usually as topical application, are recommended either when the yeasts have been isolated or in the absence of a favorable response to the previous interventions [14,44]. Additionally, denture deficiencies and other risk factors should be identified and addressed.
Denture stomatitis – clinical aspect (a); thermography of the oral mucosa (b); thermography of the maxillary denture (c)
Edentulousness and dentures can lead to muscle changes, which are mainly an adaptation to the anatomical and functional changes. These can be encountered to the muscle that define the extension of the denture base and the neutral zone and play a role in the denture stability and retention (lips, cheeks and tongue), to masticatory muscles and to the muscles of facial expression.
Prosthetic treatment deficiencies favor abnormal muscular changes. Increased vertical dimension of occlusion and ill-fitting dentures cause muscle spasms, habitual and involuntary movements. Oversized anterior buccal flange of the maxillary denture associates the overextension of the upper lip, with possible anatomical and functional consequences. Association of posterior artificial tooth wear with over jet or lack of coincidence of maximal intercuspal position and centric relation leads to an abnormal protruded mandibular position, which makes difficult the registration of maxillomandibular relationship (centric relation).
The changes in muscle tonus can be seen as hypertonia or hypotonia. Muscle hypertonia (Figure 7) is more obvious in lower lip orbicularis oris muscle and in tongue muscles, and causes instability of the mandibular denture. It occurs in the edentulous patients in relation to prosthetic factors as ill-fitting dentures, to patient’s individual characteristics as hypodivergent skeletal class II pattern, to parafunctions as bruxism or some systemic conditions. Muscle hypotonia is more frequent for upper lip orbicularis oris muscle and the buccinator muscle, and it occurs related to ageing, to deficient nutritional status and various systemic conditions. Less favorable condition for denture retention and stability, decrease of the efficiency of self-cleaning and reduced visibility of the anterior maxillary teeth in phonation or smiling are some of the effects of muscle hypotonia.
Lower lip orbicularis oris muscle hypertonia, that affects mandibular denture stability\\
The changes in volume of the muscles is usually represented by muscular atrophy, which combined with muscular hypotonia, lead to the characteristic facial aspect of old people, with masseter muscle thickness and loose or sagging skin.
Buccinators, orbicularis oris and tongue muscles define the neutral zone, whose accurate limitation is difficult to identify in severe ridge resorption. Changes in the position of the muscle insertions occur, such as high muscle insertions, even on the ridge top (genioglossus and mentalis muscle), with detached oral mucosa. Considering that position of muscle attachments has a major impact to denture base stability and retention, through changes of the denture bearing area, severe ridge resorption with consecutive muscles changes increase the treatment difficulty degree, especially in the mandible.
Muscle force decreasing leads to decrease in the capacity of performing a voluntary act (such as mastication). This occurs in relation to ageing, paresis, depression, denture instability or pain caused by the dentures. Alterations in jaw movements can occur in relation to deficiencies of the prosthetic restorations, as unstable occlusion, denture instability, increased vertical dimension of occlusion or in bruxism. Muscular spasms are encountered in particular situations as in the jaw-closing muscles, related to an increased vertical dimension of occlusion or for jaw-opening muscles related to a decreased vertical dimension of occlusion.
Neuromuscular coordination and control deficiencies, which occur in relation to age and systemic alterations, can increase treatment difficulty and negatively influence the accommodation with the prosthesis. For example, in Parkinson disease a lack of neuromuscular coordination occurs, which leads to difficulties in registration of maxillomandibular relationship and in the insertion and removal of the denture or the overdenture. Abnormal, involuntary, patterned or stereotyped and purposeless orofacial movements (oral dyskinesia) can occur linked to ill-fitting unstable dentures, oral discomfort, and lack of sensory contacts [2]. Facial nerve paresis includes affected unilateral facial musculature movement with asymmetry of facial expression and functional disorders, taste alterations and salivary changes, all having impact on the prosthetic treatment – difficulties in impression taking and in registration of maxillomandibular relationship, reduced masticatory efficiency with unilateral mastication, increased risk of unstable dentures, aesthetic alterations and denture intolerance.
If muscle changes have been identified, these should be taken into account in planning and performing the prosthodontic treatment. In muscle hypertonia, aspects like positioning the artificial teeth in the neutral zone, correct placement of the occlusal plane and correct occlusal relations are essential. In muscle hypotonia, it is recommended to design the buccal flange of the denture with a convex shape and usage of medium viscosity impression materials, in order to have a correct registration of the extension of the denture base and to use the muscle contractions for denture stabilization. Impression taking technique varies according to case’s particularities – in patients with protruded tongue at rest, wider movement are required during impression taking, comparing to a retracted tongue, in order to adequately register functional movements (Figure 8).
Tongue position at rest – anterior vs. posterior
Extension of denture or overdenture base is limited by the muscle insertions, their encroachment causing, during muscle contraction, movement of the prosthesis. In severe ridge resorption cases, as for those with muscle insertions on the ridge top, preprosthetic surgery for repositioning of muscle and mucosal attachments is indicated [51].
In neuromuscular coordination and control deficiencies, considering the severe functional alterations, conventional dentures usually don’t respond to patient’s need and implant overdenture should be chosen instead. Compared to conventional dentures, implant overdentures provides better functional parameters – exertion of higher masticatory forces promotes better nutrition through the ability to chew harder foods.
Last but not least, manufacturing of a new prosthesis requires an adjustment period for the establishment of the new memory patterns for the masticatory muscles, of about 6 to 8 weeks, aspect that should be mentioned to the patient [52].
The complete edentulism contributes greatly to the facial aspect known as the aged appearance. Prosthetic treatment needs to adequately address this consequence of edentulism, considering the fact that patients’ complaints are frequently related to aesthetic reasons.
Edentulism associates significant anatomical and functional changes that impact the facial appearance. Lip and cheek support is severely altered by tooth loss and bone resorption. A tendency of increasing the facial concavity occurs in relation to the different pattern of bone resorption of the jaws (centripetal in the maxilla and centrifugal in the mandible). In association with the loss of the occlusal contacts, a counter-clockwise rotation of the mandible, with a decreasing height of the lower third of the face, and sometimes a tendency to a more advanced protruded mandibular position occurs. Facial alterations that are directly linked to edentulism can be considered worsening factors of the esthetic appearance, since there are also preexistent changes in relation to other factors.
As a consequence of aging, there are changes related to the evolution of bones and soft tissues (muscles, fat and skin), in addition to noticeable effects of gravity, with effect on facial esthetics [53]. Systemic health, medication use and behavior (e.g., alcohol and tobacco use) can influence the facial appearance. For example, smoking causes changes particularly in the lower and middle third of the face, like hyperpigmentation and accentuated wrinkles-deeper nasolabial folds, upper lip wrinkles, lower lip vermillion wrinkles, lower lid hyperpigmentation [54]. Premature aged appearance occurs in some diseases like Cutis laxa or glomerulonephritis [55,56].
The prosthetic treatment of the edentulous patient addresses positively some of the previous mentioned facial alteration, but can also contribute to an aged appearance through its deficiencies, as in cases with a decreased vertical dimension of occlusion, a reverse smile line or darker, yellow artificial teeth.
Facial appearance of edentulous patient, with severe bone resorption, without dentures
In edentulous patient, shape and vertical proportions of the face are modified compared to the dentate period. Frequently, edentulous patients have a short face morphotype, appeared in relation to the decrease in the facial lower and total height and the counter-clockwise rotation of the mandible.
Profile changes occur as decreasing its convexity compared to the dentate period. This aspect is due to the different pattern of bone resorption of the jaws and sometimes an advanced protruded mandibular position in the absence of stable occlusion. These changes are more obvious in the skeletal class III patients and are termed as pseudo-class III relation or the old man\'s prognathism. Profile changes include also modification of nasolabial angle related to nose tip lowering and loss of upper lip support.
Lips register great changes, as reduction of vermilion height and their volume, color modifications, retraction due to support loss, elongation (upper lip) and shortening (lower lip), straight or reversed lip line and low smile line, and reduced lips dynamics that contribute to a decreased teeth exposure during speaking and smiling, which associated a reduction of emotional display, as happiness or sadness [57].
Facial changes related to ageing mark the facial appearance. Lips and cheeks become less prominent and there can be noticed marked folds and wrinkles, loose or sagging skin, changes in the skin texture and hyperpigmentation. These are mainly connected to muscle changes, as hypotonia, and skin changes, as loss of skin elastic recoil.
The prosthetic treatment has a positive impact on the facial esthetics (Figure 10). Generally, it provides a support for the soft tissue, tries to compensate the tooth loss and bone resorption (through the artificial teeth and anterior buccal maxillary flange), ensures a functional vertical dimension of occlusion and give a natural look through exposure of the teeth during smiling or speaking. Some faulty prosthesis or some changes that occurs in time can have a negative impact on facial esthetic. Unpleasant facial appearance can be linked to errors in anterior artificial tooth mounting (too forward, too backward), shade selection (chosen incorrectly, too light, not matching the patient\'s age), to changes of the artificial teeth over time (through teeth wear the smile line can become reversed, or through aging of the material discolorations can appear). A decreased vertical dimension of occlusion leads to an aged appearance, with deeper perioral folds, and an increased vertical dimension of occlusion associate an unnatural, tensioned look. An overextended buccal flange, encountered more often in the maxillary dentures, leads to an over-supported lip with a tensioned unnatural look. Unstable dentures negatively influence facial appearance through movement while speaking and the facial changes related to protruded mandibular position that many times is associated.
Facial appearance of a recently edentulous patient with and without the dentures
Removable dental prosthesis are described as having a series of complications in relation to the correctness and accuracy of their planning and execution (extension of the denture base, registration of maxillomandibular relationship, mounting of the artificial teeth, occlusal scheme), the technical and biomechanical features of the devices, the properties of the materials used, in conjunction with their evolution in time.
Considering the aims of medical treatments, not properly achieving the prosthodontic treatment goals (denture retention and stability, patient’s satisfaction that is liked to aspects like the degree of esthetic and functional rehabilitations and absence of pain) may be considered treatment complications. Removable prosthesis instability can be caused by incorrect denture execution (e.g., overextended flanges, incorrect mounting of the artificial teeth, unstable occlusion), or can occur in time, as a consequence of bone resorption. This issue must be promptly addressed since it can lead to serious complications, such as the fracture of the prosthesis, abutment loss (teeth, implants) and intolerance of the prosthesis. In order to ensure good removable prosthesis stability, the primary aspect that should be consider is its correct execution, mainly regarding the extension of the denture base and artificial teeth mounting. Secondary, usage of denture adhesives, relinings and placement of dental implants should be considered.
The fracture of the removable prosthesis (Figure 11) is a relatively common complication, having numerous risk factors, such as poor denture design, denture instability, teeth or fixed restorations in the opposite jaw, increased mucosal resiliency, previous fractures, accidents (dropping the denture, associated to reduced dexterity), material properties and changes in time, flexural fatigue or other impact factors. Its management includes identifying the cause and the treatment can range from conventionally repairing procedures to reinforcement of the denture base with metal or non-metal products (as glass and polyethylene fibers or net), to changing the previous denture or even the treatment option [58].
Overdenture fracture at the attachment site
The complications associated to the properties of the material used, mainly polymethylmethacrylate (PMMA), are linked to changes that appears during their evolution in time, as discolorations, artificial teeth wear, increased porosity and decrease flexural strength. Considering their functional and aesthetic impact, denture and overdenture treatment should be renewed at approximately every 5 years.
Additionally, signs of combination syndrome can appear when mandibular overdentures (supported or retained by roots or dental implants) are opposed by an edentulous maxilla. In this situation the masticatory field moves anteriorly, favoring the instability of the maxillary denture and the increased bone resorption rate in the anterior maxilla. This iatrogenic effect can be managed by using implants also in the maxilla, aiming to address or prevent this functional consequence and the destructive process of the oral structures [59].
The root overdentures can have teeth related complications, mainly due to primary or recurrent caries, periradicular lesions developed by vital teeth, endodontically lesions developed by endodontically treated teeth due to loss of the restoration sealing the root canal, periodontitis or root fracture [60]. Their management is dependent of the problem type, in most severe forms tooth loss and recurrent failure of prosthodontic treatment occurring. It is important to preserve the roots as a prevention factor for bone resorption and due their positive impact on the oral functioning [61]. Patients’ awareness, instruction and motivation regarding maintaining a proper oral hygiene are essential considering that is the main factor for periodontal disease and caries control. When caries occur, it is important to identify them quickly in order to have high a high success rate for the treatment. Topical fluoridation or coverage with metallic caps can be performed preventively for patients with a high caries risk. For the periodontal disease it is recommended to use Chlorhexidine 0.12% mouthwash twice daily. Also, the removal of the denture overnight and maintenance of proper denture hygiene are recommended. If tooth mobility appears, it can be addressed by reducing the tooth height, which leads to an increase in the crown to root ratio. The risk of root fracture is higher in endodontically treated teeth and when the magnitude of occlusal forces is higher, as in denture instability, bruxism, increased vertical dimension of occlusion, when teeth or fixed prosthesis in the opposite jaw. Preventively, thimble crowns can be used.
For the implant overdenture, the implants complications can be related to the treatment planning (insufficient implant number), implant positioning (surgical complications can appear, such as nerve or blood vessel injuries, penetration of the maxillary sinus or the nasal cavity, hemorrhages or pain) and their evolution (post-insertion infections, compromised survival or implant loss associated deficient osseointegration, peri-implantitis, implant fracture) [62].
Peri-implant soft tissue lesions-clinical aspect
Therefore, treatment planning considering the fundamental principles of removable implant prosthodontics, overdenture design and execution, maintenance procedures, regular check-ups are all essential for prevention or adequate management of treatment complications. Implant problems are differently addressed according to their type and severity, ranging from simple denture adjustments and enhancing the oral hygiene, to denture relinings or replacement of the denture, to inserting new implants. An important aspect to consider is that implant failure is more common in the maxilla than the mandible, consequently being favorable to place more implants in the upper jaw.
Mandibular implant overdenture is generally considered as being a good predictable treatment, its major implant complication, namely implant loss usually occurring in the first year of function [63,64]. Therefore, regular check-ups are absolutely necessary in this period, for an early intervention that ensures the best prognosis. It is recommended that the dentists performs periodically an accurate evaluation of the implants and surrounding soft tissue regarding the peri-implant marginal bone loss, implant mobility, peri-implant soft tissue, peri-implant bleeding, implant sensitivity during function, result of implant percussion test, plaque accumulation. The overdentures must be verified regarding the overdenture base that is in direct contact with the implant, as risk factor for peri-implant soft tissue complications, regarding the occlusion and maxillomandibular relationship whose faults may be related to exerting increased pressure on implants, as risk factor for implant failure, as its stability and hygiene. Other aspects, like the prosthetic treatment on the opposite jaw (an unstable denture as antagonist can produce excessive forces on the implants) and parafunctions should be checked.
Attachment system complications can occur as a consequence of an incorrect treatment planning, improper treatment conduct (e.g., errors during placement of the retentive housing in the overdenture base) or related to their changes that occur in time, during functioning (e.g., loosening or damage). These vary according to the type of attachment system, e.g., bar, ball, Locator. Most frequent attachment system complications, with overdentures, are: decreased prosthesis retention due to deactivation, detachment, damage or loss of the retentive housing; abutment screw loosening or fracture; fracture of the attachment system components (e.g., bar or clip fracture); soft tissue lesions as hyperplasia under the bar or peri-implant mucositis.
The management of attachment system complications varies according to the attachment system used and the complication type. Technical complications are more common for bar than ball attachments, and both of them are more common compared to locator system [65,66]. Usually low severity complications occurs, such as loss of rubber ring and matrix deactivation, which need to be promptly addressed since they cause overdenture instability with possible negative impact on the dental implants. A more severe complication is bar fracture, that requires increased clinical time and expenses to be resolved, considering that usually the overdenture must be replaced. In elderly edentulous patients simpler prosthetic reconstructions, with complications that require decreased time and money are preferred. Thus, if the option of implant overdenture has been selected, the ball attachment system can be more appropriate than the bar attachment system, due to the more simple maintenance procedures and easier replacement of the implant if necessary.
The conventional dentures are the most common treatment option for the edentulous patients, and usually register good results in terms of patient’s satisfaction. Dissatisfaction reasons most claimed by patients are related to denture instability, improper mastication, esthetic deficiency and phonation problems [67]. Denture intolerance is usually connected to subjective factors (the patient’s needs and expectations, psychological type, misconceptions) or objective factors (denture instability, pain, functional deficiencies).
The root or implant overdenture have improved retention that contributes to physical and psychological comfort. According to the current evidence, mandibular implant overdentures provide a higher satisfaction and oral health related quality of life compared to conventional denture, but there is uncertainty about the true magnitude of difference between the two [68].
Dentures and overdentures, the most frequently used treatment options for the complete edentulism, have complications that are related to patient and prostheses features. Patient’s general and local conditions and behavior must be acknowledged as their manifestations, interactions and impact on the prosthetic treatment. Removable implant prosthodontics principles should be well-known and respected during prosthesis execution. The previous, additional to regular check-ups, represent the basis of the prevention removable prosthesis complications.
Denture and overdenture complications are partially similar, differences being related to design particularities, biomechanical aspects and execution procedures. Addressing them depends on their nature and severity, requiring a specific medical conduct. Often simple clinical interventions are needed, but sometimes complex procedures with increased clinical, biological and financial costs must be considered in order to achieve a medical result that corresponds to the current medical standards and patient needs and expectations.
Phloem is the vascular plant tissue responsible for the transport and distribution of sugars produced by the photosynthesis. Since the plant is a continuum, phloem will be found in the external part of root cylinders (Figure 1a), in the stem vascular bundles (Figure 1b) and in the abaxial part of the venations of every single leaf (Figure 1c). While the most common is to have the phloem external to the xylem in roots and stems and abaxial in leaves, some exceptions exist and are usually taxon specific. The phloem found in the inside is named internal or intraxylary phloem (Figure 1b).
Location of the primary phloem in different organs and its cell composition. (a)
As a constitutive tissue in the plant body, phloem functions extrapolate its main function of sugar transport, including transport of signalizing molecules such as mRNAs, hormones, defenses from biotic and abiotic agents, sustenance of the organs, gas exchange, and storage of many ergastic materials, such as starch, calcium oxalate crystals, and tannins. Parenchymatic cells of the phloem can also give rise to new meristems, such as the phellogen or cork cambium. All vascular plants have phloem, which typically includes specialized living conducting cells named sieve elements whose nucleus, ribosomes, and other organelles degenerate during maturation, making sugar transport more efficient. The life and function of these cells will then rely on closely associated parenchyma cells which support the physiological functions of these sieve elements [1]. Although typical phloem is exclusive of vascular plants, rudimentary phloem-like conducting cells are present also in other lineages, such as the bryophyte leptoids, and even outside the plant kingdom, as the trumpet cells of the kelps and phaeophycean algae [2]. The primary phloem derives from the embryo and the apical meristem procambium throughout the life of the plant or from the cambium, in plants with secondary growth.
The phloem is a complex tissue and is formed typically by three cell types, the sieve elements, the parenchyma cells, and the sclerenchyma cells (Figure 2a–d). Sclerenchyma cells might sometimes be absent in primary and/or secondary phloem. The presence, quantities, and arrangements of these cell types in the tissue commonly vary and may be taxonomic informative [3, 4]. Lists depicting these variations in all phloem cell types are of ultimate importance for complete bark descriptions [5]. What follows is a description of these three major cell types in the phloem.
General aspects of the secondary phloem. (a) Composition of the secondary phloem of
Sieve element is a general term that encompasses all conducting cells of the phloem, both sieve cells and sieve tube elements [1, 6]. The name sieve derives from the strainer appearance given to the cells by the presence of numerous pores crossing their bodies (Figure 2c). These pores are specialized plasmodesmata of wider diameter, and the sieve areas are basically specialized primary pit fields [7]. The sieve pores are usually lined up with callose, which were shown to be related with the formation of the sieve pores in angiosperms, although not in gymnosperms [8]. Large amounts of callose deposit in the sieve areas also when the sieve element loses conductivity, suffers injury, or becomes dormant. Callose in gymnosperms is typically wound callose [8]. Callose can be easily detected with aniline blue under fluorescence or resorcin blue [9] (Figure 2b and c).
Sieve elements have only primary walls, but sometimes this wall can be very thick receiving the name of nacreous walls (Figure 2d) [10] and can be present in all major vascular plant lineages [1]. Nacreous walls can be very thick, and some authors have proposed they would be secondary walls [1, 8]. Nacreous walls can almost occlude the entire lumen of the sieve element (Figure 2d); hence, its presence needs to be considered in experiments of sugar translocation. Such thick walls might be related to resistance to high turgor pressures within the sieve elements. Nacreous walls seem to have a strong phylogenetic signal and are much more common in some families, such as
There are basically two types of sieve elements: sieve cells and sieve tube elements. The sieve tube elements are distinguished by the presence of sieve plates, that is, sieve areas with wider and more abundant sieve pores, usually in both extreme ends of the cells, while sieve cells lack sieve plates [1, 6, 8]. A group of connected sieve tube elements form a sieve tube [8]. According to this concept, lycophytes and ferns have sieve cells [1]. However, because of the many differences in the morphology and distribution of protoplasm organelles and chemical substances between the sieve elements of gymnosperms and vascular cryptogams, Evert [8] suggests the use of “sieve cell” as exclusive to the gymnosperms, leaving the more general term “sieve element” to the lycophytes and ferns.
The longevity of sieve elements varies. In many species it is functional for just one growth season, while for other species they can be functional a couple of years, or in the case of plants that lack secondary growth, they will be living for the entire plant life spam. Palm trees would perhaps be the plants with the oldest conducting sieve tube elements, since some reach 200 years [11]. In other plants, on the other hand, the sieve elements collapse a few cells away from the vascular cambium, corresponding to a fraction of the mm. In a mature tree, most of the secondary phloem will generally be composed of sieve elements no longer conducting. This region is called nonconducting phloem, in opposition to the area where sieve elements are turgid and conducting, called conducting phloem [5, 8] (Figure 2e and f). The term collapsed and noncollapsed phloem and functional and nonfunctional phloem are not recommended, since in some plants the nonconducting phloem keeps its sieve elements intact (Figure 2f), and although large parts of the phloem may not be conducting, the tissue as a whole is certainly still functioning in storage, protection, and even dividing or giving rise to new meristems, such as the phellogen and the dilatation meristem of some rays [5, 8].
Sieve cells are typically very elongated cells with tapering ends (Figure 3b), which lack sieve plates, that is, lack an area in the sieve element where the pores are of a wider diameter. Even though the sieve areas may be more abundant in the terminal parts of the sieve cells, the pores in these terminal areas are of the same diameter as those of the lateral areas of the sieve element. Sieve cells lack P-protein in all stages of development. The sustenance of the sieve cells is carried by specialized parenchyma cells in close contact with the sieve elements, with numerous plasmodesmata, which maintain the physiological functioning of the sieve cells, including the loading and unloading of photosynthates. These cells are known either as albuminous cells or Strasburger cells. The name albuminous was initially coined given the proteinaceous appearance of these cell’s contents. However, because the high protein content is not always present, the name Strasburger cell, paying tribute to its discoverer Erns Strasburger, is recommended over albuminous cells [5, 12]. Strasburger cells in the secondary phloem can be either axial parenchyma cells, as is common in
The secondary phloem of conifers. (a) Transverse section of the secondary phloem of
A synapomorphy of the angiosperms is the presence of sieve tube elements and companion cells, both sister cells derived from the asymmetrical division of a single mother cell. In some instances, these mother cells can divide many times, creating assemblages of sieve tube elements and parenchyma cells ontogenetically related [15]. Sieve tube elements have specialized areas in the terminal parts of the sieve elements in which a sieve plate is present (Figures 2b and c). Within the sieve plate, the pores are much wider than those of the lateral sieve areas, evidencing a specialization of these areas for conduction [16]. In
Even in lineages of angiosperms where vessels were lost and tracheids re-evolved, such as
Sieve tube elements vary morphologically. The sieve plates can be transverse to slightly inclined (Figure 2b) or very inclined (Figure 2c) and contain a single sieve area (Figure 2b) or many (Figure 2c). When one sieve area is present, the sieve plate is named simple sieve plate, while when two to many are present, the sieve plates are called compound sieve plates. Compound sieve plates typically occur in sieve tube elements with inclined to very inclined sieve plates (Figure 2c). In addition, sieve elements with compound sieve plates are typically longer than those with simple sieve plates. Evolution to sieve elements of both sieve area types has been recorded in certain lineages, such as in
In the primary phloem, just one type of parenchyma is present and typically intermingles with the sieve elements (Figure 1d). In the secondary structure, there are two types of parenchyma: axial parenchyma and ray parenchyma (Figures 2b, c, 3b, c), derived, respectively, from the fusiform and ray initials of the cambium.
The axial parenchyma in conifers commonly is arranged in concentric, alternating layers (Figure 3a and b). These parenchyma cells contain a lot of phenolic substances, which were viewed as a defense mechanism against bark attackers [21]. In Gnetales, the phloem axial parenchyma appears to be intermingling with the sieve cells (Figure 4a) [22]. Some of these axial parenchyma cells act as Strasburger cells [13].
Phloem axial parenchyma distribution in secondary phloem. (a)
In angiosperms, the distribution of the axial phloem parenchyma is more varied, and it may appear as a background tissue where other cells are dispersed or may be in bands (Figure 4b and c) and radial rows or sieve-tube-centric (Figure 4d) [5, 20]. The distribution of axial phloem parenchyma is commonly related to the abundance of fibers or sclereids. In species with more fibers, it is common to have a more organized arrangement of the parenchyma. For example, in
Although collectively described and referred to as axial phloem parenchyma, it is important to note that in many plants there will be distinct groups of phloem parenchyma within the phloem with quite different ergastic contents and therefore presumed different functions. Some of these specialized parenchyma cells may be considered secretory structures. Within a single plant, it is not uncommon that while some cells have crystals (especially when in contact with sclerenchyma), others have tannins, starch, and other substances. In apple trees (
Within bands of axial parenchyma, canals with a clear epithelium may be formed in many plant groups such as
While the phloem ages and moves away from the cambium, its structure dramatically change, and typically axial parenchyma cells enlarge (Figures 4a and b, 6c), divide, and store more ergastic contents toward the nonconducting phloem. In plants with low fiber content, the dilatation undergone by the parenchyma cells typically provokes the collapse of the sieve elements. The axial parenchyma in the nonconducting phloem can dedifferentiate and give rise to new lateral meristems. In plants with multiple periderms, typically new phellogens are formed within the secondary phloem, compacting within the multiple periderms large quantities of dead, suberized phloem. In plants with variant secondary growth, especially lianas, new cambia might differentiate from axial phloem parenchyma cells [24]. In the Asian
Sclerenchymatic cells are those with thick secondary walls, commonly lignified. Sclerenchyma can be present or not in the phloem, and when present it typically gives structure to the tissue. For instance, a phloem with concentric layers of sclerenchyma cells is called stratified (Figures 2e, 3a, and 4c) [5]—not to be confused with storied, regarding the organization of the elements in tangential section. In Leguminosae, bands of phloem are associated to the concentric fiber bands (Figure 4c).
Older phloem shows more sclerification than younger phloem, and the sclerenchyma may also act as a barrier to bark attackers [21]. The sclerenchyma is typically divided in two categories: fibers and sclereids. These cell types differ mainly in form and size, but origin has also been used to distinguish them [26].
Fibers are long and slender cells, derived from meristems, the fiber primordia [1, 26, 27]. In the primary phloem, fiber caps are sometimes found in association with the protophloem (Figure 5a) and are named protophloem fibers. Since only an ontogenetic study can evidence whether these fibers indeed differentiate within the protophloem, a term coined in the nineteenth century German and American literature, pericyclic fibers, has been recommended to be used instead of primary phloem fibers or perivascular fibers [5]. In the monocotyledons, fibers are commonly an important component of the vascular bundles (Figure 5b–d). Commonly these fibers are associated with the phloem (Figure 5b), but they might also be associated with the xylem (Figure 5c) or be central in the vascular bundle (Figure 5d). These fibers are not, however, understood as part of either phloem or xylem; although they are of vascular nature, they differentiate directly from procambium.
Vascular fibers associated to eudicot and monocot primary structure. (a) Pericyclic fiber cap (fc) and primary phloem (pp) in
Sclereids may have different forms and sizes (Figure 6a–c). Within the phloem, they are more typically square or polygonal (stone cells) and contain numerous pits and conspicuous pit canals. Holdheid [26] defines that a sclereid is a cell derived from the belated sclerification of a parenchyma cell, and that is in fact the rule in the majority of cases (Figure 6a and b). However, there are lineages in which the sclereids differentiate very close to the cambium (e.g.,
Sclereids in the secondary phloem. (a) Sclereids (sc) differentiate from parenchyma cells (arrow) in the nonconducting phloem of
On the other hand, there are cases where long and slender cells derive from previously mature parenchyma cells and are morphologically difficult to distinguish from fibers. In these cases, these cells are called fiber sclereids and may be even in concentric layers, such as in apple trees and pears (
The rays in the conducting phloem have typically the same organization in terms of width, height, and cellular composition as the secondary xylem. In this respect the rays vary from uniseriate to multiseriate (Figure 7a) and may be homocellular or heterocellular (Figure 7b). Homocellular rays are those composed of cells of one shape, all procumbent or all upright (common in many shrubs). Heterocellular rays are those where more than one cell shape is present together (Figure 7b). Ray composition is appreciated in radial sections.
Rays in the secondary phloem. (a) Longitudinal tangential section of
Because the vascular cambium produces much more xylem to the inside than phloem to the outside, phloem rays typically greatly dilate toward the periphery of the organ (Figure 7c). It is not uncommon that a dilatation meristem longitudinal to the cambium forms in some barks (Figure 7c), especially in families with very wide, wedge-like rays such as the
Rays are typically exclusively parenchymatic; however, in many species sieve elements appear in the rays and are called ray sieve cells or radial sieve cells [5, 28, 29]. These cells were recorded connecting two different sieve tubes (collections of sieve tube elements). Ray sieve elements seem to be present in taxa where perforated ray cells have been also recorded [30].
The primary phloem derives from the embryo in the seed and the procambium from the organ’s apices. Similarly to the primary xylem, the primary phloem is divided in protophloem and metaphloem (Figure 1d), with the protophloem differentiating first, while the plant is still elongating, and the metaphloem differentiating last. The phloem is always exarch, independently of the organ. Protophloem sieve elements sometimes lack companion cells, such as in
The primary phloem is simpler than the secondary phloem and is basically formed by sieve elements and parenchyma cells (Figure 1a–d). Fiber caps are commonly present, and they might be phloematic (Figure 5a). For a discussion on their origin, check the section on fibers above. The position of the phloem is typically external or abaxial to the xylem, but in some lineages the bundles are bicollateral (Figure 1b), and phloem is present both inside and outside (abaxial and adaxial), while in amphivasal bundles, the xylem encircles the phloem (Figure 5d), as in the secondary vascular tissues of some
Being derived from the cambium, the secondary phloem will share a number of characteristics with the secondary xylem. For instance, it is divided in an axial and radial system. The axial system is composed of sieve elements, axial parenchyma cells, and fibers, and the radial system is formed by rays, which are typically parenchymatic (Figure 2a–c). Similar to secondary xylem, the secondary phloem can be storied (Figure 7a) or non-storied (Figure 2b and c), depending whether the cambial mother cells are organized in tiers or not.
Some trees will have growth rings, with an early and a late phloem, both in temperate and tropical regions, but their characterization is only possible with periodical collections [5]. Sometimes, but not always, the fiber band width gives a hint on the presence of growth rings or the formation of very small sieve elements in the late phloem [1, 5].
In conifers (except Gnetales) the secondary phloem is typically marked by an alternation of axial cell types (Figure 3a and b), uniseriate rays, and, in many lineages, axial and radial resin canals (e.g.,
In other gymnosperms, in particular in Gnetales and Cycads, the first remarkable difference is the presence of very wide, multiseriate rays alternating with uniseriate rays. The wide rays in both groups have, however, evolved independently, since Cycads are a sister to all other gymnosperms, while Gnetales are within the conifers, as sister to the Pinaceae [31, 37]. In
Within the Gnetales, in
Within the angiosperms, the diversity of phloem cell type arrangements reaches its maximum. The structure can be storied (Figure 7a) or non-storied (Figure 2b and c); sclerenchyma can be present or lacking. The rays may be uni-, bi-, or multiseriate. A large array of secretory cells may be encountered, such as resin canals, laticifers, and mucilaginous cells. Crystalliferous parenchyma is also very common, especially when associated with fibers.
The variation in cell type arrangements can be of taxonomic interest. Sieve elements can vary in morphology and arrangement. They can be solitary (Figure 2f), scattered in the phloem (e.g.,
The presence, type, and arrangements of fibers and sclereids are one of the most informative characters in the bark [4]. In
Phloem parenchyma more commonly constitute the background tissue in the phloem but can also be distributed in bands (Figure 4b and c), radial rows, or even only around the sieve tube elements (Figure 4d) [5].
The classic theory of phloem transport is that proposed by Ernst Münch [42], and it involves the formation of an osmotic pressure transport gradient, where certain zones act as sources of sugars (leaves and storage organs), while others act as sinks. Experiments showed that the concentration gradients were always seen to be positive in the direction of flow [43], supporting Münch’s postulate. In a system where transport goes against the direction of transpiration, its functionality relies on the presence of a plasma membrane across the entire system to create an osmotic pressure, hence the need of a conducting system with living cells [44]. Recent studies have been refining aspects involved in the photosynthate conduction to explain long-distance transports across large trees with such a simple system [44, 45]. A direct role of intracellular calcium has also been reported in the dissolution of nondispersive P-proteins and facilitation of transport [46]. Likely, the anatomical structure of the phloem discussed in the previous sections of this chapter will prove to play a role in the system. For instance, phloem sieve element length scale with the tree sizes and sieve plate type [45]. It was also shown that sieve element’s diameter, length, and pore width increase from the top to the base of the trees [47, 48].
Across the entire pathway, sugars are removed from the system to sustain all cells in the plant body. This mechanism is only possible with the concerted mechanism between sieve elements and their close related cells (Strasburger cells and companion cells), with these accompanying cells constantly channeling substances and macromolecules toward the sieve elements [44]. The Strasburger and companion cells carry the loading and unloading of the sieve elements. Given the function of loading and unloading, the companion cell-sieve tube element size ratio is directly related to being in the source or the sink of sugars [44]. For instance, in leaves the companion cells are typically much larger, for they have the high demand of constantly loading the sieve tubes. In areas of release of the sugars (unloading), the companion cells are much smaller or even absent [44].
In the economic uses, it is not always easy to distinguish the use of the phloem from that of the periderm, since both together compose the bark of a woody plant. The phloem corresponds to the inner bark, and the periderm to the outer bark. The bark has a long history of utilization, from the production of remedies [49], aphrodisiacs (yohimbe), insecticides [50], dyes, tannins [50], angostura, fibers [51], gums and resins [50], latex, and flavorings [52].
In indigenous groups from British Columbia (Canada) and Tanzania, barks from dozens of species of woody plants are used as carbohydrate food, medicine, fibers, and structural material [50, 53]. In Mexico the bark of
The rubber tree,
I would like to express gratitude to Ray F. Evert, Veronica Angyalossy, Carmen Marcati, and André C. Lima for allowing their slide collections to be photographed and Leyde N. Nunes for the photo of
The authors declare no conflict of interest.
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Gobas",authors:[{id:"53467",title:"Dr.",name:"Juan Jose",middleName:null,surname:"Alava",slug:"juan-jose-alava",fullName:"Juan Jose Alava"},{id:"156374",title:"Dr.",name:"Frank",middleName:null,surname:"Gobas",slug:"frank-gobas",fullName:"Frank Gobas"}]},{id:"80211",title:"Perspective Chapter: Status of Dolphin in the Maritime Area of Bangladesh",slug:"perspective-chapter-status-of-dolphin-in-the-maritime-area-of-bangladesh",totalDownloads:85,totalCrossrefCites:0,totalDimensionsCites:0,abstract:"The aquatic mammalian species is the best indicator for the health of water specially dolphins. Due to human anthropogenic activities, dolphin survival rate and movement are reduced. The dreadful conditions of coastal habitats can have major concerns for dolphin population and distribution. Some dolphins swim in a short distance and some swim in a long distance. Southeast Asia is a significant geographical region for dolphin conservation. Several dolphins are highly important for the maritime area of Bangladesh that were sighted in the coastal and marine water such as Irrawaddy dolphin, Indo-Pacific Humpback dolphin, Bottlenose dolphin, Spotted dolphin, Spinner dolphin, and Ganges dolphins. Marine protected area (MPA) is a valuable zone for dolphin conservation as well as biological species. This primary information of dolphins helps for further investigation in the Bangladeshi water. The research action plan must be considered with coastal habitat, marine protected area and fishing community to conserve dolphins. We should be concerned about dolphin conservation through local and international community to develop the environment and the blue economy. Local community directly involved in the maritime area due to livelihood opportunities.",book:{id:"11335",slug:null,title:"Marine Mammals",fullTitle:"Marine Mammals"},signatures:"Md. Muzammel Hossain",authors:null},{id:"40763",title:"When Whales Became Mammals: The Scientific Journey of Cetaceans From Fish to Mammals in the History of Science",slug:"when-whales-became-mammals-the-scientific-journey-of-cetaceans-from-fish-to-mammals-in-the-history-o",totalDownloads:3495,totalCrossrefCites:3,totalDimensionsCites:5,abstract:null,book:{id:"2363",slug:"new-approaches-to-the-study-of-marine-mammals",title:"New Approaches to the Study of Marine Mammals",fullTitle:"New Approaches to the Study of Marine Mammals"},signatures:"Aldemaro Romero",authors:[{id:"139025",title:"Dr.",name:"Aldemaro",middleName:null,surname:"Romero",slug:"aldemaro-romero",fullName:"Aldemaro Romero"}]},{id:"40768",title:"Uruguayan Pinnipeds (Arctocephalus australis and Otaria flavescens): Evidence of Influenza Virus and Mycobacterium pinnipedii Infections",slug:"uruguayan-pinnipeds-arctocephalus-australis-and-otaria-flavescens-evidence-of-influenza-virus-and-my",totalDownloads:2748,totalCrossrefCites:6,totalDimensionsCites:12,abstract:null,book:{id:"2363",slug:"new-approaches-to-the-study-of-marine-mammals",title:"New Approaches to the Study of Marine Mammals",fullTitle:"New Approaches to the Study of Marine Mammals"},signatures:"Juan Arbiza, Andrea Blanc, Miguel Castro-Ramos, Helena Katz, Alberto Ponce de León and Mario Clara",authors:[{id:"142157",title:"PhD.",name:"Juan",middleName:null,surname:"Arbiza",slug:"juan-arbiza",fullName:"Juan Arbiza"},{id:"142185",title:"BSc.",name:"Alberto",middleName:null,surname:"Ponce De Leon",slug:"alberto-ponce-de-leon",fullName:"Alberto Ponce De Leon"},{id:"142190",title:"MSc.",name:"Andrea",middleName:null,surname:"Blanc",slug:"andrea-blanc",fullName:"Andrea Blanc"},{id:"142219",title:"Dr.",name:"Miguel",middleName:null,surname:"Castro-Ramos",slug:"miguel-castro-ramos",fullName:"Miguel Castro-Ramos"},{id:"142221",title:"Dr.",name:"Helena",middleName:null,surname:"Katz",slug:"helena-katz",fullName:"Helena Katz"},{id:"142222",title:"Dr.",name:"Mario",middleName:null,surname:"Clara",slug:"mario-clara",fullName:"Mario Clara"}]},{id:"40764",title:"Host-Virus Specificity of the Morbillivirus Receptor, SLAM, in Marine Mammals: Risk Assessment of Infection Based on Three-Dimensional Models",slug:"host-virus-specificity-of-the-morbillivirus-receptor-slam-in-marine-mammals-risk-assessment-of-infec",totalDownloads:2577,totalCrossrefCites:1,totalDimensionsCites:4,abstract:null,book:{id:"2363",slug:"new-approaches-to-the-study-of-marine-mammals",title:"New Approaches to the Study of Marine Mammals",fullTitle:"New Approaches to the Study of Marine Mammals"},signatures:"Kazue Ohishi, Rintaro Suzuki and Tadashi Maruyama",authors:[{id:"127528",title:"Dr.",name:"Tadashi",middleName:null,surname:"Maruyama",slug:"tadashi-maruyama",fullName:"Tadashi Maruyama"},{id:"139905",title:"Dr.",name:"Kazue",middleName:null,surname:"Ohishi",slug:"kazue-ohishi",fullName:"Kazue Ohishi"},{id:"141969",title:"Dr.",name:"Rintaro",middleName:null,surname:"Suzuki",slug:"rintaro-suzuki",fullName:"Rintaro Suzuki"}]}],onlineFirstChaptersFilter:{topicId:"301",limit:6,offset:0},onlineFirstChaptersCollection:[{id:"81844",title:"Typical Changes in Carbon and Nitrogen Stable Isotope Ratios and Mercury Concentration during the Lactation of Marine Mammals",slug:"typical-changes-in-carbon-and-nitrogen-stable-isotope-ratios-and-mercury-concentration-during-the-la",totalDownloads:4,totalDimensionsCites:0,doi:"10.5772/intechopen.103067",abstract:"The increase and decrease in the δ15N values of offspring owing to the suckling of δ15N-enriched milk (nursing) and the feeding shift from milk to solid food (weaning), respectively, are thought to be common traits observed in mammals. However, there are a few studies on lactation in marine mammals, especially large whales, because samples of calf, lactating mother, and milk are difficult to obtain. In this chapter, we review the studies on reproduction of marine mammals using δ13C and δ15N values analyzed in several tissues and describe the typical changes reported to date in those values and Hg concentrations in offspring and milk during lactation. Next, we present data on ontogenetic changes in δ15N and δ13C profiles and Hg concentration, especially focusing on the lactation period, in muscle samples of hunted bowhead whale, and stranded common minke whale (mysticetes), Dall’s porpoise (odontocete), and the harbor seal (phocid). Finally, we compare the δ15N and δ13C values in muscle samples of calves from common mink whale, Dall’s porpoise, and killer whale and suggest that these values could be excellent proxies for maternal forging habits and trophic levels.",book:{id:"11335",title:"Marine Mammals",coverURL:"https://cdn.intechopen.com/books/images_new/11335.jpg"},signatures:"Tetsuya Endo and Mari Kobayashi"},{id:"81209",title:"Phylogeny and Population Genetic Structure of Minke Whales Worldwide: A Review of Recent Studies",slug:"phylogeny-and-population-genetic-structure-of-minke-whales-worldwide-a-review-of-recent-studies",totalDownloads:28,totalDimensionsCites:0,doi:"10.5772/intechopen.102675",abstract:"In 1998, two species of minke whales were recognized based on the review of the morphological and genetic information available at that time: the Antarctic minke whale (Balaenoptera bonaerensis), which is restricted to the Southern Hemisphere, and the cosmopolitan common minke whale (Balaenoptera acutorostrata). Furthermore, three sub-species of the common minke whale were recognized: the North Atlantic (B. a. acutorostrata), North Pacific (B. a. scammoni) and Southern Hemisphere (B. a. subsp.). This chapter reviews the genetic studies on minke whales conducted after 1998. The review is organized by topic, e.g., those studies focused on phylogeny and other matters most relevant for taxonomy, and those focused on population genetic structure within oceanic basins most relevant for conservation and management. On the former topic, the new genetic information, whilst strongly supporting the minke whale taxonomic classification recognized in 1998, also reveals substantial genetic differentiation within the Southern Hemisphere common minke whales, with subsequent taxonomic implications. On the latter topic, results from different analytical procedures have provided information on population identification and structure in the Indo-Pacific sector of the Antarctic and western North Pacific, but they have failed to identify unequivocally any population within the North Atlantic common minke whales.",book:{id:"11335",title:"Marine Mammals",coverURL:"https://cdn.intechopen.com/books/images_new/11335.jpg"},signatures:"Luis A. Pastene, Mutsuo Goto, Mioko Taguchi and Yoshihiro Fujise"},{id:"81556",title:"Marine Mammals in Syria",slug:"marine-mammals-in-syria",totalDownloads:34,totalDimensionsCites:0,doi:"10.5772/intechopen.104475",abstract:"The Syrian marine water is one of the least studied areas for cetaceans in the Mediterranean Sea. Lack of basic knowledge, such as species composition and habitat, makes it impossible to develop effective conservation measures. The survey carried out along the Syrian coast by monitoring the stranding individuals on the shore since 2002 showed that there were 11 species of marine mammals living in/or visiting the Syrian marine waters at present of which 10 species belonging to the cetacean order and on belonging to pinnipeds order. The following species have been recorded: Pseudorca crassidens, Megaptera novaeangliae, Physeter macrocephalus, Tursiops truncatus, Stenella coeruleoalba, Delphinus delphis, Ziphius cavirostris, Grampus griseus, Balaenoptera physalus, Balaenoptera acutorostrata, and Manchus manchus. On the other hand, there are four species whose presence in the Syrian marine waters was mentioned a century ago by the researcher Gruvel and his team during three missions (1929–1931), but neither alive nor dead have been seen in this area during the surveys that were carried out since 1996 until the present, these species are: Phocoena phocoena, Globicephalus melas, Phocoena communis, Hyperoodon rostratus, Balaenoptera musculus. These observations reflect the vulnerability of marine mammals to anthropogenic activities, such as fishing operations, shipping, seismic activities, and climate change.",book:{id:"11335",title:"Marine Mammals",coverURL:"https://cdn.intechopen.com/books/images_new/11335.jpg"},signatures:"Adib Saad and Ilene Mahfoud"},{id:"81288",title:"How Do Whales See?",slug:"how-do-whales-see",totalDownloads:41,totalDimensionsCites:0,doi:"10.5772/intechopen.104564",abstract:"The eyes of two whales Balaenoptera physalus and Baleoptera borealis were studied by our group. In this chapter, we present the anatomical, histological, immunohistochemical and ultrastructural studies of the eyes of both types of whales. Based on the results, we can conclude that at least in these two species, the whales are rod monochromat; their resolution is very limited due to the reduced number of retinal ganglion cells, some of which were giant size (more than 100 micrometers in diameter). The excellent representation of melanopsinic positive retinal ganglion cells suggests an adaptation to the dim light as well as involvement in the circadian rhythms. The large cavernous body located in the back of the eye may provide a mechanism that allows them to move the eye forward and backwords; this may facilitate focusing and provide protection from cold deep-sea temperatures.",book:{id:"11335",title:"Marine Mammals",coverURL:"https://cdn.intechopen.com/books/images_new/11335.jpg"},signatures:"Elena Vecino, Xandra Pereiro, Noelia Ruzafa and Sansar C. 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They provide a basis for understanding dolphins’ physiology and a means for monitoring health conditions as well as furthering ecotoxicology studies.",book:{id:"11335",title:"Marine Mammals",coverURL:"https://cdn.intechopen.com/books/images_new/11335.jpg"},signatures:"Reyna Cristina Collí-Dulá and Ixchel Mariel Ruiz-Hernández"},{id:"80211",title:"Perspective Chapter: Status of Dolphin in the Maritime Area of Bangladesh",slug:"perspective-chapter-status-of-dolphin-in-the-maritime-area-of-bangladesh",totalDownloads:86,totalDimensionsCites:0,doi:"10.5772/intechopen.102022",abstract:"The aquatic mammalian species is the best indicator for the health of water specially dolphins. Due to human anthropogenic activities, dolphin survival rate and movement are reduced. The dreadful conditions of coastal habitats can have major concerns for dolphin population and distribution. Some dolphins swim in a short distance and some swim in a long distance. Southeast Asia is a significant geographical region for dolphin conservation. Several dolphins are highly important for the maritime area of Bangladesh that were sighted in the coastal and marine water such as Irrawaddy dolphin, Indo-Pacific Humpback dolphin, Bottlenose dolphin, Spotted dolphin, Spinner dolphin, and Ganges dolphins. Marine protected area (MPA) is a valuable zone for dolphin conservation as well as biological species. This primary information of dolphins helps for further investigation in the Bangladeshi water. The research action plan must be considered with coastal habitat, marine protected area and fishing community to conserve dolphins. We should be concerned about dolphin conservation through local and international community to develop the environment and the blue economy. Local community directly involved in the maritime area due to livelihood opportunities.",book:{id:"11335",title:"Marine Mammals",coverURL:"https://cdn.intechopen.com/books/images_new/11335.jpg"},signatures:"Md. 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He completed a one-year Post-Doctoral Fellowship awarded by the DFAIT (Foreign Affairs and International Trade Canada) at the Institute of Biomedical Engineering of the University of New Brunswick (Canada) in 2010. Currently, he is Professor in the Faculty of Electrical Engineering (UFU). He has authored and co-authored more than 200 peer-reviewed publications in Biomedical Engineering. He has been a researcher of The National Council for Scientific and Technological Development (CNPq-Brazil) since 2009. He has served as an ad-hoc consultant for CNPq, CAPES (Coordination for the Improvement of Higher Education Personnel), FINEP (Brazilian Innovation Agency), and other funding bodies on several occasions. He was the Secretary of the Brazilian Society of Biomedical Engineering (SBEB) from 2015 to 2016, President of SBEB (2017-2018) and Vice-President of SBEB (2019-2020). 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Professor Derbel has published many articles in journals and collaborates intensively with IntechOpen Access Publisher as an editor.",institutionString:"Clinique les Oliviers",institution:null},{id:"300144",title:"Dr.",name:"Meriem",middleName:null,surname:"Braiki",slug:"meriem-braiki",fullName:"Meriem Braiki",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/300144/images/system/300144.jpg",biography:"Dr. Meriem Braiki is a specialist in pediatric surgeon from Tunisia. She was born in 1985. She received her medical degree from the University of Medicine at Sousse, Tunisia. She achieved her surgical residency training periods in Pediatric Surgery departments at University Hospitals in Monastir, Tunis and France.\r\nShe is currently working at the Pediatric surgery department, Sidi Bouzid Hospital, Tunisia. Her hospital activities are mostly concerned with laparoscopic, parietal, urological and digestive surgery. She has published several articles in diffrent journals.",institutionString:"Sidi Bouzid Regional Hospital",institution:null},{id:"229481",title:"Dr.",name:"Erika M.",middleName:"Martins",surname:"de Carvalho",slug:"erika-m.-de-carvalho",fullName:"Erika M. de Carvalho",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/229481/images/6397_n.jpg",biography:null,institutionString:null,institution:{name:"Oswaldo Cruz Foundation",country:{name:"Brazil"}}},{id:"186537",title:"Prof.",name:"Tonay",middleName:null,surname:"Inceboz",slug:"tonay-inceboz",fullName:"Tonay Inceboz",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/186537/images/system/186537.jfif",biography:"I was graduated from Ege University of Medical Faculty (Turkey) in 1988 and completed his Med. PhD degree in Medical Parasitology at the same university. I became an Associate Professor in 2008 and Professor in 2014. I am currently working as a Professor at the Department of Medical Parasitology at Dokuz Eylul University, Izmir, Turkey.\n\nI have given many lectures, presentations in different academic meetings. I have more than 60 articles in peer-reviewed journals, 18 book chapters, 1 book editorship.\n\nMy research interests are Echinococcus granulosus, Echinococcus multilocularis (diagnosis, life cycle, in vitro and in vivo cultivation), and Trichomonas vaginalis (diagnosis, PCR, and in vitro cultivation).",institutionString:"Dokuz Eylül University",institution:{name:"Dokuz Eylül University",country:{name:"Turkey"}}},{id:"71812",title:"Prof.",name:"Hanem Fathy",middleName:"Fathy",surname:"Khater",slug:"hanem-fathy-khater",fullName:"Hanem Fathy Khater",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/71812/images/1167_n.jpg",biography:"Prof. Khater is a Professor of Parasitology at Benha University, Egypt. She studied for her doctoral degree, at the Department of Entomology, College of Agriculture, Food and Natural Resources, University of Missouri, Columbia, USA. She has completed her Ph.D. degrees in Parasitology in Egypt, from where she got the award for “the best scientific Ph.D. dissertation”. She worked at the School of Biological Sciences, Bristol, England, the UK in controlling insects of medical and veterinary importance as a grant from Newton Mosharafa, the British Council. Her research is focused on searching of pesticides against mosquitoes, house flies, lice, green bottle fly, camel nasal botfly, soft and hard ticks, mites, and the diamondback moth as well as control of several parasites using safe and natural materials to avoid drug resistances and environmental contamination.",institutionString:null,institution:{name:"Banha University",country:{name:"Egypt"}}},{id:"99780",title:"Prof.",name:"Omolade",middleName:"Olayinka",surname:"Okwa",slug:"omolade-okwa",fullName:"Omolade Okwa",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/99780/images/system/99780.jpg",biography:"Omolade Olayinka Okwa is presently a Professor of Parasitology at Lagos State University, Nigeria. She has a PhD in Parasitology (1997), an MSc in Cellular Parasitology (1992), and a BSc (Hons) Zoology (1990) all from the University of Ibadan, Nigeria. She teaches parasitology at the undergraduate and postgraduate levels. She was a recipient of a Commonwealth fellowship supported by British Council tenable at the Centre for Entomology and Parasitology (CAEP), Keele University, United Kingdom between 2004 and 2005. She was awarded an Honorary Visiting Research Fellow at the same university from 2005 to 2007. \nShe has been an external examiner to the Department of Veterinary Microbiology and Parasitology, University of Ibadan, MSc programme between 2010 and 2012. She is a member of the Nigerian Society of Experimental Biology (NISEB), Parasitology and Public Health Society of Nigeria (PPSN), Science Association of Nigeria (SAN), Zoological Society of Nigeria (ZSN), and is Vice Chairperson of the Organisation of Women in Science (OWSG), LASU chapter. She served as Head of Department of Zoology and Environmental Biology, Lagos State University from 2007 to 2010 and 2014 to 2016. She is a reviewer for several local and international journals such as Unilag Journal of Science, Libyan Journal of Medicine, Journal of Medicine and Medical Sciences, and Annual Research and Review in Science. \nShe has authored 45 scientific research publications in local and international journals, 8 scientific reviews, 4 books, and 3 book chapters, which includes the books “Malaria Parasites” and “Malaria” which are IntechOpen access publications.",institutionString:"Lagos State University",institution:{name:"Lagos State University",country:{name:"Nigeria"}}},{id:"273100",title:"Dr.",name:"Vijay",middleName:null,surname:"Gayam",slug:"vijay-gayam",fullName:"Vijay Gayam",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/273100/images/system/273100.jpeg",biography:"Dr. Vijay Bhaskar Reddy Gayam is currently practicing as an internist at Interfaith Medical Center in Brooklyn, New York, USA. He is also a Clinical Assistant Professor at the SUNY Downstate University Hospital and Adjunct Professor of Medicine at the American University of Antigua. He is a holder of an M.B.B.S. degree bestowed to him by Osmania Medical College and received his M.D. at Interfaith Medical Center. His career goals thus far have heavily focused on direct patient care, medical education, and clinical research. He currently serves in two leadership capacities; Assistant Program Director of Medicine at Interfaith Medical Center and as a Councilor for the American\r\nFederation for Medical Research. As a true academician and researcher, he has more than 50 papers indexed in international peer-reviewed journals. He has also presented numerous papers in multiple national and international scientific conferences. His areas of research interest include general internal medicine, gastroenterology and hepatology. He serves as an editor, editorial board member and reviewer for multiple international journals. His research on Hepatitis C has been very successful and has led to multiple research awards, including the 'Equity in Prevention and Treatment Award” from the New York Department of Health Viral Hepatitis Symposium (2018) and the 'Presidential Poster Award” awarded to him by the American College of Gastroenterology (2018). He was also awarded 'Outstanding Clinician in General Medicine” by Venus International Foundation for his extensive research expertise and services, perform over and above the standard expected in the advancement of healthcare, patient safety and quality of care.",institutionString:"Interfaith Medical Center",institution:{name:"Interfaith Medical Center",country:{name:"United States of America"}}},{id:"93517",title:"Dr.",name:"Clement",middleName:"Adebajo",surname:"Meseko",slug:"clement-meseko",fullName:"Clement Meseko",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/93517/images/system/93517.jpg",biography:"Dr. Clement Meseko obtained DVM and PhD degree in Veterinary Medicine and Virology respectively. He has worked for over 20 years in both private and public sectors including the academia, contributing to knowledge and control of infectious disease. Through the application of epidemiological skill, classical and molecular virological skills, he investigates viruses of economic and public health importance for the mitigation of the negative impact on people, animal and the environment in the context of Onehealth. \r\nDr. Meseko’s field experience on animal and zoonotic diseases and pathogen dynamics at the human-animal interface over the years shaped his carrier in research and scientific inquiries. He has been part of the investigation of Highly Pathogenic Avian Influenza incursions in sub Saharan Africa and monitors swine Influenza (Pandemic influenza Virus) agro-ecology and potential for interspecies transmission. He has authored and reviewed a number of journal articles and book chapters.",institutionString:"National Veterinary Research Institute",institution:{name:"National Veterinary Research Institute",country:{name:"Nigeria"}}},{id:"158026",title:"Prof.",name:"Shailendra K.",middleName:null,surname:"Saxena",slug:"shailendra-k.-saxena",fullName:"Shailendra K. Saxena",position:null,profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002bRET3QAO/Profile_Picture_2022-05-10T10:10:26.jpeg",biography:"Professor Dr. Shailendra K. Saxena is a vice dean and professor at King George's Medical University, Lucknow, India. His research interests involve understanding the molecular mechanisms of host defense during human viral infections and developing new predictive, preventive, and therapeutic strategies for them using Japanese encephalitis virus (JEV), HIV, and emerging viruses as a model via stem cell and cell culture technologies. His research work has been published in various high-impact factor journals (Science, PNAS, Nature Medicine) with a high number of citations. He has received many awards and honors in India and abroad including various Young Scientist Awards, BBSRC India Partnering Award, and Dr. JC Bose National Award of Department of Biotechnology, Min. of Science and Technology, Govt. of India. Dr. Saxena is a fellow of various international societies/academies including the Royal College of Pathologists, United Kingdom; Royal Society of Medicine, London; Royal Society of Biology, United Kingdom; Royal Society of Chemistry, London; and Academy of Translational Medicine Professionals, Austria. He was named a Global Leader in Science by The Scientist. He is also an international opinion leader/expert in vaccination for Japanese encephalitis by IPIC (UK).",institutionString:"King George's Medical University",institution:{name:"King George's Medical University",country:{name:"India"}}},{id:"94928",title:"Dr.",name:"Takuo",middleName:null,surname:"Mizukami",slug:"takuo-mizukami",fullName:"Takuo Mizukami",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/94928/images/6402_n.jpg",biography:null,institutionString:null,institution:{name:"National Institute of Infectious Diseases",country:{name:"Japan"}}},{id:"233433",title:"Dr.",name:"Yulia",middleName:null,surname:"Desheva",slug:"yulia-desheva",fullName:"Yulia Desheva",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/233433/images/system/233433.png",biography:"Dr. Yulia Desheva is a leading researcher at the Institute of Experimental Medicine, St. Petersburg, Russia. She is a professor in the Stomatology Faculty, St. Petersburg State University. She has expertise in the development and evaluation of a wide range of live mucosal vaccines against influenza and bacterial complications. Her research interests include immunity against influenza and COVID-19 and the development of immunization schemes for high-risk individuals.",institutionString:'Federal State Budgetary Scientific Institution "Institute of Experimental Medicine"',institution:null},{id:"238958",title:"Mr.",name:"Atamjit",middleName:null,surname:"Singh",slug:"atamjit-singh",fullName:"Atamjit Singh",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/238958/images/6575_n.jpg",biography:null,institutionString:null,institution:null},{id:"252058",title:"M.Sc.",name:"Juan",middleName:null,surname:"Sulca",slug:"juan-sulca",fullName:"Juan Sulca",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/252058/images/12834_n.jpg",biography:null,institutionString:null,institution:null},{id:"191392",title:"Dr.",name:"Marimuthu",middleName:null,surname:"Govindarajan",slug:"marimuthu-govindarajan",fullName:"Marimuthu Govindarajan",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/191392/images/5828_n.jpg",biography:"Dr. M. Govindarajan completed his BSc degree in Zoology at Government Arts College (Autonomous), Kumbakonam, and MSc, MPhil, and PhD degrees at Annamalai University, Annamalai Nagar, Tamil Nadu, India. He is serving as an assistant professor at the Department of Zoology, Annamalai University. His research interests include isolation, identification, and characterization of biologically active molecules from plants and microbes. He has identified more than 20 pure compounds with high mosquitocidal activity and also conducted high-quality research on photochemistry and nanosynthesis. He has published more than 150 studies in journals with impact factor and 2 books in Lambert Academic Publishing, Germany. He serves as an editorial board member in various national and international scientific journals.",institutionString:null,institution:null},{id:"274660",title:"Dr.",name:"Damodar",middleName:null,surname:"Paudel",slug:"damodar-paudel",fullName:"Damodar Paudel",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/274660/images/8176_n.jpg",biography:"I am DrDamodar Paudel,currently working as consultant Physician in Nepal police Hospital.",institutionString:null,institution:null},{id:"241562",title:"Dr.",name:"Melvin",middleName:null,surname:"Sanicas",slug:"melvin-sanicas",fullName:"Melvin Sanicas",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/241562/images/6699_n.jpg",biography:null,institutionString:null,institution:null},{id:"117248",title:"Dr.",name:"Andrew",middleName:null,surname:"Macnab",slug:"andrew-macnab",fullName:"Andrew Macnab",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"University of British Columbia",country:{name:"Canada"}}},{id:"322007",title:"Dr.",name:"Maria Elizbeth",middleName:null,surname:"Alvarez-Sánchez",slug:"maria-elizbeth-alvarez-sanchez",fullName:"Maria Elizbeth Alvarez-Sánchez",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Universidad Autónoma de la Ciudad de México",country:{name:"Mexico"}}},{id:"337443",title:"Dr.",name:"Juan",middleName:null,surname:"A. Gonzalez-Sanchez",slug:"juan-a.-gonzalez-sanchez",fullName:"Juan A. Gonzalez-Sanchez",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"University of Puerto Rico System",country:{name:"United States of America"}}},{id:"337446",title:"Dr.",name:"Maria",middleName:null,surname:"Zavala-Colon",slug:"maria-zavala-colon",fullName:"Maria Zavala-Colon",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"University of Puerto Rico, Medical Sciences Campus",country:{name:"United States of America"}}}]}},subseries:{item:{id:"93",type:"subseries",title:"Inclusivity and Social Equity",keywords:"Social Contract, SDG, Human Rights, Inclusiveness, Equity, Democracy, Personal Learning, Collaboration, Glocalization",scope:"