Studies done so far to evaluate the effect of calf sex in Holstein cows’ milk production.
\r\n\t
",isbn:null,printIsbn:"979-953-307-X-X",pdfIsbn:null,doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!1,hash:"dc069c307f12ff3436be837ed7d2252a",bookSignature:"Dr. Selcan Karakuş",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/10264.jpg",keywords:"Click Chemistry, Nanostructures, Microstructures, Controllable Click Synthesis, Nanoparticle, Nanocomposite, Drug Delivery Nanosystems, Release Kinetic, Bionanoconjugation, Environmental Applications, Removal of Pollutants, Sensor",numberOfDownloads:null,numberOfWosCitations:0,numberOfCrossrefCitations:null,numberOfDimensionsCitations:null,numberOfTotalCitations:null,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"July 8th 2020",dateEndSecondStepPublish:"July 29th 2020",dateEndThirdStepPublish:"September 27th 2020",dateEndFourthStepPublish:"December 16th 2020",dateEndFifthStepPublish:"February 14th 2021",remainingDaysToSecondStep:"6 months",secondStepPassed:!0,currentStepOfPublishingProcess:5,editedByType:null,kuFlag:!1,biosketch:"Dr. Karakuş worked as a visiting researcher at the Department of Polymer Science and Engineering, University of Massachusetts, USA where she gained research experience in molecular adsorption, self-assembled polymeric nanostructures and nanocomposites, and copolymer blends.",coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"206110",title:"Dr.",name:"Selcan",middleName:null,surname:"Karakuş",slug:"selcan-karakus",fullName:"Selcan Karakuş",profilePictureURL:"https://mts.intechopen.com/storage/users/206110/images/system/206110.jpeg",biography:"Assistant professor Selcan Karakus is currently working at the Department of Chemistry, Istanbul University - Cerrahpasa(IUC), Turkey. She received her Master of Science degree in Physical Chemistry from Istanbul University (IU) in 2006. Dr. Karakus received her Doctor of Philosophy degree in Physical Chemistry from IU in 2011. She has worked as a visiting researcher at the University of Massachusetts, Department of Polymer Science and Engineering. She has research experience in molecular adsorption, self-assembled polymeric nanostructures and nanocomposites, and copolymer blends. 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Cancer",doi:"10.5772/intechopen.87091",slug:"the-mechanisms-of-proliferation-and-energy-metabolism-in-oral-cancer",body:'\nCancer development and fatality are gradually increasing worldwide. It is thought that the reasons are complex but associated with both aging and increase in the population [1, 2]. With the rapid global population increase and aging, the rising significance of cancer as a leading cause of death is partly correlated with a marked decline in mortality rates due to stroke and coronary heart disease in many countries. In addition, tobacco and alcohol consumption are known to be etiologically associated with carcinogenesis. Cancer is both a genetic and progressively systemic disease. In all types of cancer, somatic cells begin to divide uncontrollably and spread into surrounding tissues. Cancer cells can arise almost anywhere in the human body. Normal cells grow and divide according to the body’s needs, and when they age or become damaged, they die and are replaced by new cells. However, when cancer arises, this orderly process breaks down: as cellular abnormalities increase, old or damaged cells survive rather than being removed, and new cells form when they are not needed. These extra cells can divide uncontrollably and may form malignant tumors. Although many cancers form solid tumors composed of masses of tissue, cancers of the blood, such as leukemias, generally do not form solid tumors [2, 3, 4]. Head and neck cancer is a common neoplasm that encompasses epithelial malignant tumors of the nasopharynx, larynx, and mouth, representing about 6% of all cases and accounting for an estimated over 650,000 new cancer cases and over 350,000 cancer-related deaths worldwide every year [5, 6]. Oral cancer is the most notably frequent cancer type in the head and neck region, squamous cell carcinoma being the most common single entity. Oral cancers comprise two categories: those affecting the oral cavity (lips, inner lips, cheeks, teeth, gums, the anterior two-thirds of the tongue, the floor, and palate) and those affecting the oropharynx (middle region of the throat, including the tonsils and base of the tongue). Such cancers may arise in any location, although there are certain areas that are affected more frequently, such as the tongue and gingiva. These areas represent about 90% of all malignancies of the oral cavity [7, 8]. However, despite significant advances in surgery and chemotherapy over the last few decades [9], oral cancer is still characterized by a poor prognosis and a low survival rate [10]. The 5-year survival rate of those diagnosed is ~ 60%.
\nIn patients diagnosed with oral cancers at an advanced stage, there is a high incidence of metastasis to surrounding tissues, lymph nodes, and distant organs [5]. Metastasis is recognized as a process, whereby genetic instability in the primary tumor accelerates cell heterogeneity, allowing a few metastatic clones to eventually emerge and be positively selected to disseminate cancer at a distance [11]. This is the most annihilating stage of malignancy and the leading cause of cancer-related death. In metastasis, cancer cells break away from the primary cancer, travel through the blood or lymph system, and form new metastatic tumors elsewhere in the body. Each metastatic tumor is the same type of cancer as the primary tumor [12, 13], and the cells of each resemble each other upon microscopic observation. Moreover, they usually share common molecular features, such as the presence of specific genetic changes. It is clear that only a minority of malignant cells participate in the process of metastasis, due to interaction with host tissues and the intrinsic characteristics of the cancer cells themselves; thus, metastasis may imply an escape of these cells from the hostile environment they have created, characterized by features such as hypoxia, inflammation, and immunological surveillance [14, 15, 16].
\nThis chapter details recent findings on molecular markers that are involved in the mechanisms of proliferation and energy metabolism of oral cancer and provides new perspectives on its diagnosis and treatments.
\nCAMs are typically single-pass transmembrane receptors [17] compounded of three stored domains: an intracellular domain that interacts with a transmembrane domain, an extracellular domain, and the cytoskeleton. These domains can interact in several different ways [18]. The first way is through homophilic binding, where the same CAMs bind to each other. They are also capable of heterophilic binding, which means a CAM on one cell will bind to different CAMs on another cell. The third type of binding is that between cells and substrate, where a mutual extracellular ligand binds two different CAMs. There are four major superfamilies, or groups, of CAMs: the immunoglobulin super family of cell adhesion molecules (IgCAMs), the cadherins, the integrins, and the c-type lectin-like domain proteins (CTLDs). Proteoglycans are also believed to be a class of CAMs. One classification system involves distinction between calcium-dependent and calcium-independent CAMs [19]. The Ig-superfamily CAMs and integrins are not dependent on calcium ions, whereas selectins and cadherins are calcium-dependent. In addition, integrins participate in cell-matrix interactions, while other CAM families play some important roles in cell-to-cell interactions [20].
\nNeural cell adhesion molecule (NCAM) is a family of cell surface glycoproteins playing an important role in the development of the nervous system, fasciculation, axonal outgrowth, regulation of cell migration, and branching [21]. NCAM has several isoforms derived from alternative splicing of a single gene [22, 23, 24]. In particular, the three major isoforms with molecular weights of 120, 140, and 180 kDa have similar extracellular parts but differ in the disposition of their domains, which are cytoplasmic for the two larger polypeptides [25, 26]. Furthermore, the expression of NCAM is upregulated by transforming growth factor (TGF)-β1 [27, 28, 29]. Although NCAM was initially considered to exist only in neural tissue, it has since been observed in the human kidney, lung, fetal muscle, and colon, as well as in elements of the hemopoietic system. Furthermore, it has been described that NCAM is expressed by a variety of human tumors and associated with perineural invasion by various neoplasms, such as gallbladder cancer, melanoma, bile duct cancer, and adenoid cystic carcinoma of the head and neck [30, 31, 32, 33, 34, 35]. We have also demonstrated previously that NCAM is sporadically found in the adenoid cystic carcinoma, derived from human submandibular salivary gland, in vivo [36]. NCAM is believed to mediate adhesion between cells through a calcium ion-independent homophilic (NCAM-NCAM) binding mechanism and to mediate adhesion between neurons and the extracellular matrix through heterophilic binding (NCAM to another ligand or counter-receptor) [21]. It has been described that exogenously added NCAM can inhibit the proliferation of cultured neonatal astrocytes and of astrocytes responding to a penetrating lesion in the adult rat brain, in vivo [37, 38], suggesting that these effects are mediated by homophilic binding to NCAM on the astrocyte membrane.
\nAdenoid cystic carcinoma (ACC) is a well-known and typical malignant salivary gland tumor. ACCs are biologically aggressive and can bring metastases even when many years have passed after excision of the primary tumor. Facial paralysis is especially frequent, causing perineural and/or neural invasion. We have attempted to examine the role of NCAM by investigating the effect of anti-NCAM antibody (MAb NCAM) and TGF-β1 in human salivary gland tumor cells. The expression and distribution of NCAM were also investigated in ACC tissues. We further found that apoptotic cell death was induced via a DNA damage signal through the mitochondria, inducing release of cytochrome c into the cytoplasm of salivary gland tumor cells [36]. However, MAb NCAM had no effect on human oral squamous cell carcinoma (HOSCC) cell lines, which do not express NCAM. As shown in Figure 1, these results indicate that the effect of MAb NCAM is specific to NCAM-expressing tumor cells, such as human salivary gland tumor cells; furthermore, blocking the ability of NCAM through MAb NCAM, as well as the homophilic (NCAM-NCAM) binding mechanism, rather than regulating a signaling pathway of cell proliferation, may in fact induce a negative signal such as apoptosis in human salivary gland tumor cells. In addition, homophilic (NCAM-NCAM) binding may activate multiple signaling pathways that differ among cell types. In view of the fact that NCAM expression on human salivary gland tumor cells is upregulated by TGF-β1, it can be hypothesized that a further homophilic (NCAM-NCAM) binding mechanism may be activated and that consequently the proliferative activity of HSG cells may also be upregulated by TGF-β1-mediated NCAM activity (Figure 2).
\nSchematic representation of MAb NCAM-induced apoptotic signal transduction pathways via the DNA damage signal through the mitochondria involved in apaf-1 and caspase activation in NCAM-expressing cells.
Schematic representation of TGF-β1-induced upregulation of NCAM expression and proliferative activity involved in homophilic (NCAM-NCAM) binding mechanism in NCAM-expressing cells.
On the other hand, cimetidine, the most studied histamine type-2 receptor (H2R) antagonist used clinically, is commonly prescribed to treat gastroesophageal reflux disease as well as gastric and duodenal ulcers [39]. Cimetidine has recently been shown to possess antitumor activity against gastric, kidney, and colon cancers, as well as melanomas [40, 41, 42, 43]. A recent study has suggested that this behavior of cimetidine is mediated through three different effects: a direct inhibitory effect on tumor growth by blocking the cell growth activity of histamine via activation of H2 receptors and an indirect effect involving inhibition of tumor-associated angiogenesis, an immunomodulatory effect through augmentation of the host’s immune response to tumor cells, and an inhibitory effect on cancer cell migration and adhesion to endothelial cells, thus inhibiting tumor angiogenesis and metastasis [44]. We have also examined the NCAM-associated impact of cimetidine on tumor growth and perineural/neural invasion in salivary gland tumors using an in vitro cell culture system and an in vivo nude mouse cancer model. These experiments clearly indicated that cimetidine effectively downregulated the expression of NCAM by inhibiting NF-κB transactivation, subsequently blocking salivary gland tumor cell adhesion to neural cells, and ultimately inducing apoptosis in salivary gland tumor cells, thus preventing the growth of salivary gland tumor masses in nude mice [45]. Although malignant glandular tumors are commonly known to be resistant to chemotherapy and/or radiation, the clinical application of cimetidine as an anticancer drug might provide an integral part of future therapeutic strategies against NCAM-expressing tumors such as adenoid cystic carcinoma.
\nFinally, it was suggested that NCAM might be associated with not only a cell-to-cell adhesion mechanism but also tumorigenesis, including the occurrence, development, and perineural/neural invasion of human salivary gland tumors.
\nFurther studies will be required to identify the signal transduction pathways by which treatment with cimetidine suppresses the growth of salivary gland tumors and to establish a strategy for cimetidine-based therapy for those tumors.
\nCoxsackievirus and adenovirus receptor (CAR/CXADR), a transmembrane glycoprotein, was initially characterized as a viral attachment site on the surface of epithelial cells (Figure 3) [46]. Later it was identified as a component of the tight junction (TJ) complex, an interacting partner for a number of other TJ proteins and a regulator of TJ formation [47, 48, 49, 50, 51, 52]. Furthermore, CAR is known to be a cell-cell adhesion molecule [53, 54]. In terms of function, loss of CAR has been considered to diminish intercellular adhesion, increase proliferation, and promote the migration as well as invasion of cancer cells [55, 56]. On the basis of these observations, a tumor-suppressive role of CAR in human cancers has speculated. Although it has recently been described [55, 56, 57, 58] that CAR is observed in various organs, it is still unclear whether it is expressed in oral cancer. Therefore, we examined the role of CAR in SCC in the oral cavity (data not shown). This revealed that CAR was constitutively expressed in five oral SCC cell lines. To analyze the function of CAR, we then examined the proliferative activity of SAS cells after CAR gene knockdown. However, CAR knockdown did not promote the proliferative activity of SAS cells. Although the expression level of CAR was decreased by CAR knockdown, that of NF-κB p65 (RelA) showed little change. Furthermore, SAS cell numbers were notably reduced by CAR overexpression. Finally, it was suggested that the overexpression of CAR in SAS cells led to apoptosis via activation of caspase-9. In addition, the localizations of CAR and RelA in 40 samples of HOSCC at various stages were investigated using immunohistochemistry. A positive reaction for polyclonal antibody (PAb) CAR was weakly observed on the membrane of carcinoma cells in 19 of 40 cases (47.5%) of HOSCC. The immunoreactivity for CAR further tended to fade at the invasive front of oral SCC (Figure 4). In the meantime, RelA immunoreactivity was strongly positive, particularly on the nucleus of carcinoma cells at the invasive front, in 30 of 40 cases (75%) of HOSCC. These observations suggest that CAR plays a significant role in the inhibition of oral cancer cell growth.
\nSchematic representation of coxsackievirus and adenovirus receptor on the tumor cells.
(A) The positive reaction for PAb CAR was observed on the membrane of tumor cells in 19 of 40 cases (47.5%) of SCCs. (B) NF-κB immunoreactivity were clearly detected in 30 of 40 cases (100%) in the HOSCC tissues. The immunoreactivity for CAR especially tended to fade away in the invasive front of oral SCC tissues. NF-kappaB (+)/CAR (−), 15/21 cases (71.4%).
Cytokines are composed of a large family of secreted proteins that bind to and signal through defined cell surface receptors on a wide variety of target cells, playing an important role in the maintenance of homeostasis. Furthermore, many cytokines share structural features and effects during inflammation, development, or immune responses.
\nThe concept of a control mechanism for cellular growth via regulation of apoptosis has recently been erected in a wide variety of tissue systems. Changes in the balance between cell survival and death are definite signs of emergence of various tumors. Therefore, modulation of apoptosis is required so as to maintain the homeostasis of a living organism. The expression of cytokines and their receptors in human oral cancers has attracted a great deal of interest because of their potential importance in tumor immunity. In particular, it has been described that members of the tumor necrosis factor (TNF) family, including Fas/FasL and TNF-related apoptosis-inducing ligand (TRAIL), regulate the deletion of unnecessary immune cells through induction of apoptosis [59, 60, 61]. However, despite their expression of these obvious antigens, tumor evasion by the immune system is often inefficient. It is considered that tumor cells may also evade immune attack by expressing TRAIL, Fas ligand, or other molecules that induce apoptosis in activated T cells [62].
\nTRAIL, also called APO2 ligand (APO2L), is a novel member of the TNF cytokine family that was originally characterized by its ability to induce apoptosis [59, 60]. It is recognized that at least four closely related receptors bind to TRAIL: death receptor-4 (DR4) and DR5/KILLER, which contain cytoplasmic death domains and signal apoptosis [60, 61]; decoy receptor-1 (DcR1) [61, 62, 63], which lacks a cytoplasmic tail and inhibits TRAIL function; and DcR2 [64, 65], which contains a cytoplasmic region with a truncated death domain that does not transduce the death signal [67]. TRAIL interacts with its agonistic receptors DR4 and DR5, inducing apoptosis in a wide variety of cancer cell lines derived from breast carcinoma, lung carcinoma, colon carcinoma, lymphoma, malignant melanoma, and malignant glioma [59, 60, 68, 69]. Although DR4 is expressed in many normal human tissues and cells, including spleen and peripheral blood leukocytes, TRAIL induces apoptosis in various cancer cells, but not in normal cells [70]. This may be explained by the fact that TRAIL also interacts with the antagonistic decoy receptors DcR1 and DcR2, which are expressed in normal tissues but not in cancer cells (Figure 5) [66, 67]. Neither DcR1 nor DcR2 receptors induce apoptosis, but they protect cells from TRAIL-induced apoptosis [64, 65, 66]. Until now, the biological involvement of the complex TRAIL receptor system has remained unclear, and the existing data are conflicting. Nevertheless, because of its selective cytotoxicity against tumor cells, TRAIL is regarded as a promising anticancer weapon that might be highly effective in vivo with few side effects, as it has little or no function on normal tissues.
\nSchematic representation of the death receptors and decoy receptors on the tumor cell.
TRAIL is expressed in most normal human cells and tissues, including the peripheral blood leukocytes, spleen, lung, and prostate, but not the brain [60]. However, the expression of TRAIL in human neoplasms is largely unknown. Accordingly, we have examined whether TRAIL and its receptors are expressed in HOSCC tissues or cell lines and whether these cell lines are sensitive to TRAIL-induced apoptosis [71]. This revealed that the mRNA and protein levels of TRAIL and its receptors are co-expressed in HOSCC cell lines in the absence of paracrine fratricide or autocrine suicide. Moreover, TRAIL protein was also detected in 24 of 50 samples (48%) of HOSCC tissue, and there was no correlation among the WHO grades. These findings suggest that HOSCC has the potential to escape immune surveillance by killing host T lymphocytes via DR4/TRAIL and DR5/TRAIL interactions, as suggested for FasL [72, 73, 74].
\nThe expression of decoy receptors in cancer cells is, however, a phenomenon that objects against previous reports [62, 63]. Indeed, several authors have currently described that a decoy receptor is expressed in various cancer types and our results are consistent with their findings [75, 76, 77]. Therefore, cancer cells may also avoid TRAIL-induced apoptosis by expressing a decoy receptor.
\nOn the other hand, cells differ significantly in their response to TRAIL. As contrasted to HUVEC or other oral cancer cells, only KB cells undergo significant apoptosis following exposure to recombinant human (rh)TRAIL. The reason why KB cells, despite their expression of a decoy receptor, react to rhTRAIL remains unresolved.
\nCommonly, the most proximal step in suppression of a death receptor pathway is inhibition of ligand binding. This may be acquired by lack, or the presence of decoy receptors [65, 66, 67, 68], or mutations of death receptors [78, 79]. However, it has recently been described that there is no correlation between the expression of TRAIL receptor and susceptibility to TRAIL-induced apoptosis in various cancer types [75, 76, 77]. Furthermore, the existence of antiapoptotic proteins, such as bcl-2, bcl-xL, and/or fas-like IL-1-converting enzyme (FLICE)-like inhibitory protein [80, 81], also seems to be significant, as they are resistant to death receptor-mediated apoptosis. Holistically, our results also suggest that there may be no correlation between the expression of TRAIL receptor and sensitivity to TRAIL-induced apoptosis in HOSCC cell lines and that TRAIL-resistant cells (HSC-2, HSC-3, HSC-4, and Ca9-22) may express cytoprotective proteins that block TRAIL-induced apoptosis or that the apoptotic effect of TRAIL is regulated by other mechanisms. It has also been described that TRAIL, in combination with an anticancer drug, acts cooperatively to induce apoptosis in various cancer cells that are resistant to TRAIL or chemotherapy [75, 82, 83]. This combination of TRAIL with chemotherapeutic reagents might be a useful therapeutic strategy against TRAIL-resistant cell lines such as HSC-2, HSC-3, HSC-4, and Ca9-22 used in our investigation.
\nIn fact, we are currently investigating the synergistic effects of α-mangostin and TRAIL on induction of apoptosis via the mitochondrial pathway in squamous cell carcinoma of the oral cavity [84]. To summarize, mangosteen (Garcinia mangostana) is a tree discovered in Southeast Asia, and the pericarp of its fruit has been used in folk medicine for the treatment of many human diseases. The rinds of mangosteen fruit contain a high concentration of xanthone, a type of polyphenol. One form of xanthone, α-mangostin, has been described to exhibit chemopreventive effects against chemically induced colon cancer through a decrease of c-Myc expression, suppressing tumor growth in a mouse model of mammary cancer. A recent study has proved the inhibitory effect of α-mangostin on the growth of prostate cancer. However, it is still unclear whether α-mangostin induces cell death in oral cancer. Then, the present study examined the impact of α-mangostin on HOSCC. First, we analyzed the expression of c-Myc in five HOSCC cell lines (HSC-2, HSC-3, HSC-4, Ca9-22, and SAS). The highest level of c-Myc mRNA expression was found in SAS cells and the lowest in HSC-4 cells. Therefore, SAS cells were treated with α-mangostin, which was observed to exert a weak cytocidal effect. Since α-mangostin has been described to exert synergistic effects on cancers when combined with anticancer drugs, we tried to evaluate these synergistic effects of α-mangostin in combination with TRAIL. We found that this combination induced apoptosis in SAS cells through the mitochondrial pathway via activation of caspase-3/caspase-7 and caspase-9, following the release of cytochrome c. In addition, this apoptosis was induced by S/G2/M-phase arrest. Immunoreactivity for c-Myc was revealed in the cytoplasm of cancer cells in 16 (40%) of the 40 cases of HOSCC. These data showed that the combination of α-mangostin and TRAIL may have considerable potential for the treatment of oral cancer.
\nFurther investigation of TRAIL-mediated cell death, including the interaction of TRAIL and its receptors in oral cancer cells under various conditions, will be required to establish a strategy for TRAIL-based oral cancer therapy, which does not cause liver toxicity.
\nThe tyrosine kinase receptor c-Met ordinarily binds with hepatocyte growth factor (HGF), which triggers its involvement in processes such as cell differentiation, cell growth, angiogenesis, and embryogenesis [85, 86]. However, c-Met activation is also associated with processes related to malignant transformation, such as invasion, tumor growth, angiogenesis, and metastasis [87, 88, 89, 90, 91]. In addition to autocrine or paracrine signaling via HGF, c-Met may also be activated via the mutation, protein overexpression or amplification, or transcriptional alteration (via microRNA-mediated dysregulation) of MET [92, 93, 94]. c-Met overexpression and MET amplification are thought to be associated with a poorer prognosis in some types of tumors, including non-small cell lung cancer and gastric cancer [94]. In HOSCC, it has also been reported that c-Met expression is associated with cisplatin resistance and a strong propensity for metastasis in vivo [95], as well as a poor prognosis [96]. However, details of the involvement of c-Met in oral carcinogenesis are still unclear. Accordingly, we investigated how the relationship between the expression of c-Met and several tumor activation-related markers such as NF-κB is associated with oral carcinogenesis (data not shown). In addition, the expression and distribution of c-Met and NF-κB were also examined in HOSCC tissues (data not shown). The results of real-time qRT-PCR and immunoblot analysis indicated overexpression of c-Met mRNA and protein in SAS cells. Therefore, SAS cells were used in this study. To investigate how c-Met functions in SAS cells, c-Met knockdown analysis was performed. c-Met knockdown appeared to reduce the number of SAS cells. To confirm whether this had been due to apoptosis, caspase activity was then analyzed, and this revealed that apoptosis had indeed occurred via activation of caspases-9 and caspases-3/caspases-7 in SAS by c-Met knockdown. Furthermore, SAS showed cell cycle arrest at S/G2/M phase during this apoptotic cell death. Subsequently, to determine NF-κB expression after c-Met knockdown, we also used a siRNA approach to reduce the expression of c-Met and determine the effects on NF-κB activity. As expected, the level of c-Met mRNA was markedly reduced by c-Met siRNA. Moreover, c-Met knockdown by c-Met siRNA clearly decreased the activation of NF-κB mRNA in SAS cells, in comparison with controls. These data indicated that c-Met upregulated NF-κB activation and consequently that c-Met knockdown led to apoptosis of SAS cells. These combined data suggested that c-Met produced by autologous cancer cells promoted tumor growth. Furthermore, our in vivo studies demonstrated c-Met-specific immunoreactivity, consistent with the observation of NF-κB-positive cells in HOSCC biopsy samples (c-Met expression, 9/20 (45%); NF-κB expression, 18/20 cases (90%)). This result suggests that c-Met expression correlates with increased activation of NF-κB. Based on these in vitro and in vivo observations, it can be hypothesized that c-Met function leads to NF-κB activation and subsequently anti-apoptosis and that as a consequence, it may be associated with tumorigenesis, including growth, development, and angiogenesis in HOSCC.
\nFurther investigations of the role of c-Met will be required to fully understand c-Met-mediated tumor proliferation and to establish a therapeutic strategy for c-Met-based oral cancer.
\nIn normal tissues, adenosine triphosphate (ATP) is mainly produced in mitochondria via complete oxidative phosphorylation (OXPHOS) of glucose. Conversely, only 10% of ATP is produced from glycolysis in which glucose is replaced to lactate [97]. Interestingly, cancer tissues possess high levels of glycolysis in the cytosol even under aerobic conditions, which is upregulated by PI3K/Akt signaling in the mitochondria, a phenomenon known as the “Warburg effect” or “aerobic glycolysis” [98, 99, 100, 101, 102, 103]. More than 80 years ago, Otto Warburg found that cancer cells expended much more glucose to generate lactic acid than normal cells even under conditions of O2 sufficiency. This finding was the first indication that cancer cells have an alteration of glucose metabolism. Warburg [99] considered that these defects of respiration caused a form of metabolic disturbance that was significant for carcinogenesis. After Warburg, many biologists attempted to clarify the molecular basis of aerobic glycolysis occurring in tumor cells. Accumulated evidence suggested that many cancer-related genes, such as p53, c-Myc, and Ras, are all associated with modulation of the Warburg effect [104]. As a master regulator of the cancer hypoxic response, hypoxia-inducible factor (HIF)-1 plays very important roles in modulating aerobic glycolysis to meet the biosynthetic demands of cancer cells and to protect them from damage due to hypoxic stress [105]. Warburg theorized that cancer cells shift from oxygen-dependent efficient ATP production via OXPHOS in mitochondria to the less efficient cytoplasmic glycolysis. As a result, cancer cells need to burn up more glucose to maintain their energy requirements for survival and growth. It has been reported that HIF-1α activates the expression of glucose transporter 1, 3 (Glut1, Glut3) under hypoxic conditions [106, 107], which acquires sufficient glucose uptake by tumor cells. We also examined HIF, and details can be found in Chapter 10 of the InTech book Tumor Microenvironment and Myelomonocytic Cells [108]. Although clarification of glucose metabolism is considered vital for understanding energy metabolism in oral cancer, lipid metabolism has also been receiving attention recently (Figure 6).
\nSchematic representation of the connecting glucose and lipid metabolism in cancer cells.
Lipids are composed of phospholipids, triglycerides, cholesterol esters, cholesterol, fatty acids, sphingolipids, and other molecules, which are critical components of cellular membranes [109, 110, 111, 112, 113]. In addition to their role as structural components, lipids also act as energy resources and as signaling molecules to sustain cell growth [114, 115, 116]. Lipid metabolism is known to be largely altered in cancers [117, 118, 119, 120], and worsening lipogenesis has been indicated to be a predominant characteristic of most tumors [114, 115, 121]. In oral cancer, cancer tissues include higher levels of unsaturated fatty acids than those in normal tissue (data not shown). Recent studies have revealed intrinsic molecular alterations in lipid metabolism. Especially, fatty acid synthase (FAS) is a key enzyme for synthesis of fatty acid from acetyl CoA, which is expressed at high levels in the adipose tissue and liver but at low levels in other tissues in humans [122]. FAS is overexpressed in several human cancers, including those of the ovary, bladder, stomach, breast, lung, prostate, oral cavity, and melanoma, and this overexpression is associated with poor prognosis [123, 124]. Furthermore, glutathione peroxidase 4 (GPX4) expression in tumors is positively correlated with tumor survival and linked to pathways that regulate cell proliferation, motility, and tissue remodeling [125]. Knockdown of GPX4 suppresses the formation and progression of cancer and leads to non-apoptotic cell death, ferroptosis. We have also examined the role of GPX4 in HOSCC (data not shown). Ferroptosis is a non-apoptotic form of cell death that can be triggered by conditions or small molecules that inhibit the glutathione-dependent antioxidant enzyme GPX4 or glutathione biosynthesis. This lethal process is defined by depletion of plasma membrane polyunsaturated fatty acids and the iron-dependent accumulation of lipid reactive oxygen species. It has also been reported that GPX4 is negatively regulated by the p53 gene [126]. These data suggest that GPX4 plays a significant role in proliferation and progression and may serve as a potential therapeutic target in HOSCC. Thus, GPX4 would be useful as a predictor of poor outcome in patients with oral cancer, and its antibody might be applicable as an inhibitor of oral cancer progression. Identification of the signaling pathways underlying these events might help to elucidate the mechanism of development of oral cancer. Further investigations into the role of GPX4 will be required to fully understand GPX4-mediated cancer proliferation and to establish a GPX4-based therapeutic strategy for oral cancer.
\nUnlike surgery, chemotherapy, and radiotherapy, which can have serious side effects on the human body, the use of agents that have no such side effects, such as TRAIL, cimetidine, mangostin, and antibodies, for cancer therapy mobilizes and regulates systemic functions, enhancing the body’s ability to fight cancer. Therefore, these medicines may be more appropriate for patients with inoperable advanced cancer, those in periods of chemotherapy intermission, or those during postoperative recovery. There has been an increased emphasis on such agents for prevention of cancer and inhibition of cancer metastasis. There has been an impressive renaissance in the search for semi-synthetic drugs or derivatives from natural compounds. Progress in this regard not only adds to the chemical bank but also leads to a better comprehension of the chemical basis of treatments lacking side effects for the treatment of cancers using drugs obtained from natural sources.
\nFurther studies will be required to establish a strategy for basic molecular and clinical approaches for effective oral cancer therapy, which should be tailored to individual patients.
\nWe have no financial relationships to disclose.
\nReproduction in mammalian females demands high energetic costs, driving the mobilization of fat deposits, in both gestation and lactation [1]. In evolutionary biology, numerous hypotheses defend a sex-biased allocation of these resources by the pregnant and nursing females, to maximize the reproductive success of their male and female offspring. Some of these theories support their reasoning in the local resource competition [2, 3], local resource enhancement [4], “advantaged daughters” [5], the “safe bet”/reproductive value [6, 7] and the sex-differentiated sources of mortality [8].
\nThe most well-known and tested theory remains the Trivers-Willard hypothesis that predicts that: 1) females in good body condition will allocate her offspring sex ratio towards males; 2) and that mothers in good body condition will also invest more per son than per daughter if males exhibit greater variation in reproductive value when males exhibit greater variation in reproductive value [9] According to this hypothesis, female mammals are able to adjust the sex of their offspring based on their own condition as a form to maximize reproductive success in the next generation. This theory also states that the mother will adapt her milk production to offspring gender, for example, by increasing milk production or changing its composition when she is nursing an offspring of the gender that has higher chances of producing future descendants. This strategy is particularly beneficial in species whose males compete for mating, like bovine, with dominant bulls leaving abundant offspring and weaker ones having no offspring at all. On the other side, this hypothesis also describes that investment in female offspring will be more profitable when the mothers are in poor condition because the chance of producing competitive male offspring is low. Well-nourished mothers invest more in male offspring, as strong sons will more likely leave more offspring, whereas even weaker daughters will produce more progeny than weak sons [10].
\nIn agreement with this theory it has been shown that, in humans, the milk produced for males is more energy dense in well-nourished mothers [11], while mothers with low socioeconomic status, when nursing daughters produce milk with a higher fat content than when nursing sons [12]. However, evidence for systematic sex-biased favoring males has been equivocal [13, 14, 15, 16, 17]. Post-natal, sex-biased nursing care has been investigated as a possible reason for sex-biased milk production in several mammalians, including humans. Several studies reported evidence of sex-biased milk synthesis in different species but drawing definitive conclusions from these studies has been difficult for several reasons [11, 12, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28, 29, 30].
\nThis chapter intents to discuss the evidences pro and against the existence of a sex-bias in cows´ milk production, by stressing the putative effects of the calf gender in consecutive lactations while focusing in particular in dairy cows. Albeit non-consensual, its existence would open new insights on the economic impacts of using sex sorted semen to enhance farm productivity.
\nIn the last decades, industrial intensive milk production system uses the Holstein-Friesian breed, known as highest milk producing cow in the world. It is well documented that, with almost no exceptions, there has been a continuous increase in milk yield per cow. In all countries milk production and milk composition evolved over the years, due to a higher genetic merit and better management of the cows [31, 32]. For example, from 2002 to 2013, in Denmark, all but two years showed a significant increase in the milk production compared to the previous year [33]. In São Miguel island, Azores, the same evolution was observed [34].
\nAlso, in all countries, seasonal variations in milk production and composition were observed, both in intensive [35, 36] and pasture-based systems [37]. Previous studies have also proven that milk production varies with parity. It is generally observed a progressive increase in milk production in the first three to four parities and then a progressive decrease [38, 39].
\nThe main source of income in a dairy farm is, by far, milk sale and any factor that can increase or decrease the production affect significantly the farmers income. Even though external factors like feeding, rearing and management are an important part of profitability, other factors, intrinsic to animals can have an important impact on profitability of a farm. The genetic merit [40] and sex of the calf are some of those factors. Beside the intrinsic difference in the commercial value of a female or a male offspring in a commercial farm, if the female milk production is indeed affected by the calf gender, then it could be a major factor for maximizing profits [41].
\nCalf-sex biased milk production is the capability of a cow to adapt milk production and composition to the sex of her offspring, a phenomenon well documented in diverse mammalian species [19]. The milk yield and the quality of milk produced are two important characteristics in dairy cow production and are also of great impact in beef production. Any favoring of one sex over the other in bovine offspring can lead to a great increase in the use of sex sorted semen, despite its lower conception rate [42]. In dairy cows, birth sex-ratio is biased, with more males being born, which suggests underlying mechanisms operating to favor more male offspring [43, 44].
\nBull calves in dairy farms are mostly unwanted, due to their low value; in some countries, they are euthanized after birth, raising an ethical and social concern for the industry. On the other side, the used of sexed semen has higher costs and lower fertility. The fertility of sexed semen is estimated to be 8 to 17.9% lower in heifers compared to the conventional and not advisable to use in multiparous cows [42, 45, 46]. If a specific calf sex is associated with higher milk yield, this would have obvious consequences in the value and widespread use of sexed semen [46].
\nThe growth rate of a suckling a male calf is higher than in females [47]. Therefore, it would be expected cows to have higher milk production or more energy dense milk when nursing a male. Despite differences in milk quantity or composition, cows do not show any sex biases in nursing behavior [29]. Since in most dairy farms, contrasting to most beef operations, calves are removed from the mother soon after calving, the pre and peri-natal mechanisms are the sole responsible for any observed milk-production sex-bias. Besides, cows are usually pregnant for most of the previous lactation [48], so the calf sex can potentially influence the previous lactation during its gestation or the lactation after their birth.
\nIn Bovidae, data on the effect of calf sex in milk production are, to the least, inconsistent. Some studies reported an effect of calf sex on milk yield [1, 49, 50, 51], whereas other studies found no association [52]. One of the studies found that cows with a given genotype had higher milk yield in case of a male calf than a heifer calf [51]. In buffalos no effect between calf sex and milk production was reported [53].
\nIn dairy cows in particular, studies addressing milk production sex-bias so far led to different results. While most studies described an advantage of female offspring, this effect was not observed for all the populations and a significant difference was not always observed [1, 33, 34, 40, 41, 54, 55]. Canadian and Iranian data for calf-sex bias in milk production found milk yield to be increased when a heifer was calved [41, 56]. However, a higher milk yield after calving a female offspring was only seen in the second lactation in New Zealand Holstein-Friesians [40], and only in the first lactation in French Holstein-Friesians [54] (Table 1).
\nCountry | \nYears analyzed | \nN (lactations) | \nBreed | \nMain results | \nStudy (chronologically) | \n
---|---|---|---|---|---|
USA | \n1995-1999 | \n2,390,000 | \nHolstein | \nCows favor daughters, producing significantly more milk for daughters across lactation. Effects of fetal sex interact dynamically across parities, whereby the sex of the fetus being gestated can enhance or diminish the production of milk during an established lactation. Sex of the fetus gestated on the first parity has persistent consequences for milk synthesis on the subsequent parity. Specifically, gestation of a daughter on the first parity increases milk production by, 445 kg over the first two lactations | \n[1] | \n
France | \n2000-2008 | \n8,901,000 | \nMontbéliarde and Holstein | \nThe effect of the calf sex hardly affected milk production. A small effect in favor of males was observed in parity 2 and 3 and was similar across breeds. The estimated effect of the sex of the calf in gestation on the simultaneous lactation was also very small. | \n[53] | \n
Iran | \n1992-2008 | \n402,716 | \nHolstein | \nCows with female calves had higher milk and fat yield, persistency of milk and fat yield and longer lactation length, while cows that gave birth to male calves had shorter calving interval and longer productive life. Also, cows with female calves had higher milk yield per day of lactation in the first two parities, but there was no difference in milk yield per day of lactation for parities ≥3. | \n[40] | \n
Denmark | \ntill 2013 | \n71,088 | \nHolstein | \nCows produced higher volumes of milk if they had a bull calf, with a significantly higher milk production of 0.28% in the first lactation period for cows giving birth to a bull calf. Such difference was even higher when cows gave birth to another bull calf, so having two bull calves resulted in a difference of 0.52% in milk production compared to any other combination of sex of the offspring. | \n[32] | \n
New Zealand | \n1995-2005 | \n578,912 | \nHolstein and Jersey | \nThe lactation initiated by the birth of a female calf was associated with a 0.33–1.1% (p < 0.05) higher milk yield. Female calf gender present carryover effects associated with higher milk yield in second lactations for Holstein Friesians (0.24%; p = 0.01) and third lactations for Jerseys (1.1%; p = 0.01). | \n[39] | \n
UK | \n2013-2014 | \n211,932 | \nHolstein | \nGiving birth to a heifer calf conferred a 1% milk yield advantage in first lactation heifers, whilst giving birth to a bull calf conferred a 0.5% advantage in second lactation. | \n[46] | \n
Azores, Portugal | \n2009-2017 | \n45,712 | \nHolstein | \nWhether calving a male or a female, cows roughly produced the same amount of milk and protein content. However, the content of fat was slightly higher when they calved females (P < 0.001). | \n[33] | \n
Studies done so far to evaluate the effect of calf sex in Holstein cows’ milk production.
However, Hinde et al. [1], with the largest study done so far on this topic, documented sex-biased milk production in US Holstein cattle. In his population, cows favor daughters, producing significantly more milk for daughters than for sons across lactation, suggesting that the effects of fetal sex can interact dynamically across parities. The sex of the fetus being gestated can enhance or diminish the production of milk during an established lactation. Moreover, the sex of the fetus gestated on the first parity has persistent consequences for milk synthesis on the subsequent parity. Contrastingly, Gillespie et al. [57] did not detect a significant effect of the sex of the calf being gestated on the mother milk production. Dallago et al. [55]. found only a calf-sex effect on the lactose and total solids, with an advantage to the females.
\nOn a population of 1.49 million cows from the late 90’s, primiparous cows giving birth to a female produced, on average, additional 142 kg (1.3% increase) of milk over a standardized 305-day lactation period compared with those calving a male [1]. The fetal sex on the first parity had also persistent effects on milk production during the second lactation. Calving a female on the first parity, increases milk production by 445 kg over the first two lactations, identifying a dramatic and sustained programming of mammary function by the offspring in utero. On the other side, cows calving a male son on their first parity produced less milk on their second lactation (P < 0.001), particularly if they also gestated a male calf on the second pregnancy (Table 2). According to the same study, the milk composition was similar whether the gestation produced a gestation of a son or daughter; the fat concentration was 3.61% after gestation of a daughter and 3.62% after gestation of a son; protein concentrations were the same (3.17%) [1].
\nDifferences Kg (%) | \nCalf gender combination at the first and second lactation | \n||
---|---|---|---|
Female-female vs. male-male | \nFemale-male vs. male-male | \nMale-female vs. male-male | \n|
at first lactation | \n24 (0.3) | \n7 (0.1) | \n13 (0.2) | \n
at second lactation | \n52 (0.6) | \n5 (0.1) | \n53 (0.6) | \n
Cumulative effect | \n76 (0.9) | \n12 (0.2) | \n66 (0.8) | \n
Effect of the calf gender combination at the first and second lactation (305d) according to Hinde et al. [1].
Gillespie et al. [57] also showed that, in the UK, calving a heifer was associated with a 1% milk yield advantage in first lactation heifers, but calving a bull calf conferred a 0.5% advantage in second lactation. Heifer calves were also associated with a 0.66 kg reduction in saturated fatty acid content of milk in first lactation, even though there was no significant difference between genders in the second lactation. Interestingly, the effects of calf gender observed on both the yield and saturated fatty acid content were considered minor compared to the nutritional and genetic influences. Aspects that affect milk production, such as mastitis [58] or lameness [59], seem to have a deeper impact on milk production than calf gender.
\nAn Iranian study, using 402,716 Holstein milk records from 1991 to 2008, report that cows calving a female offspring present a higher milk and fat yield and longer persistency of milk and fat yield, as well as a longer lactation length [41]. Cows calving a male offspring presented shorter calving interval and an overall longer reproductive life. The observed higher daily milk yield after calving a female in the first two parities was not maintained for the next parities [60]. However, a higher occurrence of dystocia in male calving was not taken into consideration and was most likely a factor for the higher milk production observed after calving a female calf [41]. In contrast, both a French [54] and a Danish [33] studies found a small increase in milk yield in both Holstein and Montbéliarde dams calving a male offspring. On the French study, the sex-bias favoring males effect reached 40 kg milk (0.5% of the mean), 0.6 kg fat, 0.6 kg protein. A small difference was also noticed for fat and protein contents (from 0.01 to 0.02%) in parity 2 and 3. Similarly, the estimated effect of the sex of the calf in gestation on the simultaneous lactation is very small [54].
\nGræsbøll et al. [33] also reported significantly higher milk productions (0.28%) in first lactating cows producing a bull calf. This difference was even higher when cows calved another bull calf, with a difference of 0.52% in milk production compared to any other possible combinations of offspring sex. The same study pointed that dams would favor a bull fetus by decreasing milk production during the second pregnancy if the calf born in the first parity was a heifer, which diluted the positive effect on milk production of calving a male in the first pregnancy. Being pregnant with a bull fetus may reduce milk production to possibly increase the energy spent on the bull fetus. Also, cows seem to favor living bull offspring over unborn bull offspring, but unborn bull offspring over living heifer offspring [33].
\nThe magnitude of sex bias milk production, when observed in other species, seems to be stronger among first parity females [11, 26, 27, 30, 59]. The fetal sex effect may be disguised in multiparous females because of the cumulative effects of sequential gestations with fetuses of different sexes on the mammary gland architecture [1]. It is also possible that maternal investment tactics may change according to the residual reproductive value of the offspring [1, 61] or transmit a targeted effort during a critical window of mammary gland preparation for a new lactation [1, 62]. Interaction effects were observed between calf gender across the first three parities, with the lowest second parity milk yield observed when a cow gave birth to male calves in all three parities. First parity calf sex did not have a significant effect on the third lactation milk yield. Disparities between the effects for calf sex sequences that differed only by the calf gender in the first parity were not significantly different from each other [40].
\nIn cows’ populations were a daughter-biased milk production was observed, this may involve life-history tradeoffs for both cows and their daughters. High milk production in dairy cows has been associated with reduced fertility, health, and survival depending on environmental conditions [63]. It was also observed that cows gestated during lactation have moderately reduced survival and milk production in their own adulthood [48, 64].
\nSome of the differences found across different studies could be partly explained by differences in the datasets used; Hess et al. [40] used total lactational yield, calculated using the test interval method; Hinde et al. [1] and Barbat et al. [54] used the test day model rather than predicting 305 day milk yields; Graesboll et al. [33] adopted a farm-based approach using Wilmink curves to calculate 305 day milk yields and Gillespie et al. [47] used the Milkbot lactation model, that can be affected by environment and genetics [65]. Also, the use of sexed semen was not known in most of the studies and it can have a significant impact in the results obtained. Sexed semen is mainly used to breed heifers with higher genetic merit [54, 66] and this creates an obvious bias towards female calves. This can be aggravated by the fact that heifers inseminated with sexed semen tend to have lower fertility and become pregnant later, consequently calving in an older age, which is associated with a higher milk production [67].
\nThe effect of the calf gander can further interact with other factors, like parity or seasonality, making it difficult to evaluate it in a precise way. It was observed that after the third calving, the mother milk production was independent of the calf gender. This observation might be related to larger pelvic dimensions of older cows and by consequence a lower incidence of dystocia [41].
\nA significant difference between the dairy industry in Azores [34] or New Zealand [40] compared with other populations is that both are primarily pasture-based. The production and calving in Azores are not, however, as seasonal as the one observed in New Zealand [34, 40]. In the non-seasonal pasture-based system no calf-sex bias in milk yield was observed, even though a slight increase in fat percentage was associated with the birth of a male calf [34].
\nIn Denmark, the difference observed in milk production due to the sex of the offspring was generally smaller than the difference between farms. Other management related aspects are more important for the milk yield registered and the differences identified might be due to size of the offspring rather than the sex, but size and sex might also have separated effects [33]. So far, no relation was observed among mean somatic cells count and the sex of the calf born [41, 68, 69], even though this parameter is often associated with the cow body condition [70].
\nModeling complex biological features, such as milk production, is challenging due to the number of inherent and environmental aspects that can influence them. Also, the statistical model used for analysis may influence to a certain point the results and data interpretation. One explanation for the differences of the several studies on calf-sex biased milk production can be related to the models used. For example, in one of the studies, Holstein Friesian cows calving males in the first three parities had significantly lower first lactation milk yield than cows calving two males followed by a female in the first three parities, but this observation is biased if models do not include lactation length. Also, there are no reasonable biological reasons why to test the effect of the gender of the third calf on the first lactation yield. In fact, the observed effect of calf gender on milk yield is due to an association between calf gender and milk yield rather than calf gender triggering a difference in milk yield. The alleged effect of the third parity calf gender on the first lactation milk yield was not apparent when lactation length was included in the models [40].
\nIn beef cows, studies with limited samples led to different sex-biased milk production, pointing to either favors a son [71], or a daughter [72], or not show any sex-biases [73]. A study in the red Chittagong cattle found no effect of the calf-sex in milk production [74].
\nIn New Zealand, with a seasonal calving system, the calf gender was reported to influence milk yield possibly through the increased gestation length of male calves [40]. In that study, the milk production tests were performed on the same date for all cows, so those calving a male would have their tests performed, on average, 2 days earlier. However, when the lactation length (reported longer in male calves) was included in the model, no effects existed of the calf gender over the increased production of milk [40]. At least part of the reported difference in milk production due to calf gender, was really due to methodological issues. The interval-centering method used provide a 10.8 ± 4.0 L higher milk yield if herd tests are 2 days later in lactation. However, the observed calf sex variance is too large to be explained only by this difference in herd test dates. When lactation length shortens depending on calving date, as well as the herd tests occurring 2 days earlier, the difference in milk yield is 26.9 ± 6.2 L. This difference is similar to the observed effect of calf gender on milk yield, further supporting that this effect is, at least partially, due to the different lactation length when male calves are born 2 days later [40].
\nIt is difficult to establish any association between the calf gender and a presumed sex-biased milk production or a sex-biased pregnancy length, because of various existing confounder factors that may permeate such interaction. Mean pregnancy in length male calves is longer than in females, the difference also being affect by breed and parity [40, 75]. Also, primiparous cows tend to present shorter pregnancies than multiparous cows, the calves born lighter [76], albeit the risk for dystocia is also higher for first calving cows.
\nRecently, Atashi and Asaadi [77], using 252,798 lactations on 108,077 Holstein dairy cows in Iran showed that multiparous cows with longer gestations performed better in lactation than primiparous cows. This study also showed that multiparous cows with short gestation length had a lower yield at the beginning of lactation and higher raising and declining slopes of the lactation curve compared with cows presenting longer or average length of pregnancy.
\nThe production system may also interfere with milk production performance of dairy cows. In seasonal breeding systems, late calving cows usually have a shorter lactation since the entire herd ceases lactation on the same day [40]. The lactation length is usually longer in non-seasonal systems because the lactation can continue until the milk yield of an individual cow drops below a point when it is more economical to dry the cow. In these conditions a weaker negative correlation between gestation and lactation length is observed compared with seasonal systems where all the cows are dried of on a single day [40]. However, even in non-seasonal systems lactation length was observed to be approximately four days shorter following the birth of a male calf compared to a female calf across the first four lactations [41]. Chegini et al. [41] found that cows calving female offspring had more persistent lactations than those that calving male offspring, suggesting that the lactation curves are different.
\nStill, there is some controversy regarding the best methodology to apply when modeling the milk production (whether in milk yield or composition) to adequately account the effects of the gender of the calf. This is not an easy task, because it establishes a complex interaction with other parameters (e.g., pregnancy length, dystocia, and some cow related factors) that may act as confounding factors. Lactation length is one important factor affecting milk yield per lactation, leading to the need to introduce correction factors for lactation length in the models for milk production in cows. Lactation length in itself has a negative relationship with the annualized production of milk and milk solids [78]. Also, the milk yield and milk production curves change according to the lactation number, the persistency of the peak and lactational length, the cow genetics and the number or milking frequency, among other factors. Such aspects should also be considered in the lactation modeling studies. Models construct evaluate the lactation curves should be used that take all possible confounders into account simultaneously. Therefore, further investigation is necessary to confirm whether the shape of lactation curves differ based on calf gender and identify potential biological explanations for any such difference.
\nCalving difficulty is higher with larger calves [79]. It is also known that there is a higher frequency of dystocia in male calves’ birth [60, 74, 79]. Dystocia significantly reduces the whole lactation milk yield [40, 50, 54, 69, 75, 76, 77], besides increasing veterinary treatment costs [76], and reducing cow fertility. After dystocia there is a higher incidence of metritis [77], ketosis [80, 81], both associated with a decrease in milk production. Also, an easy calving presumably leads to a higher milk production because it is associated with reduced stress and pain during calving, consequently leading to a lower energy imbalance that can cause more metabolic disorders [41].
\nMale calves are typically larger than females, and pose a greater risk of dystocia [1, 79, 82]. However, Hinde et al. [1] reported that sex-biased milk synthesis remained when analysis was restricted to a subset of females without record of dystocia, and included information on individual cows across the first and second parity, favoring females.
\nA Danish study found different results. Farmer assisted calving were associated with a higher milk yield while cows with no farmer assistance or with veterinary assistance during the most recent calving produced less milk. This means that mildly to moderate calving difficulties improved milk yield, while no assistance or the need for veterinary assistance decreased subsequent milk production. In the same study the interaction between sex of offspring and difficulty of calving was found to be insignificant [33]. Still, it must be also considered that dystocia might go unnoticed, nevertheless affecting milk production, which could lead to misreading of the sex-bias towards higher production after female calving because of unidentified or unrecorded dystocia [40]. The effect of the different degrees of dystocia in milk production or for how long they persist remain unclear [83, 84, 85].
\nIn UK Holstein-Friesian cows, moderate calving difficulties resulted in higher milk production. It is possible that some births not needing help and human supervision may experience real difficulties that go unnoticed and are wrongly registered as an easy calving, when they might have had some difficulties without the farmer’s notice. Furthermore, it is likely that cows with highly valuated genetic material may be offered calving assistance from the farmer more often [67].
\nA reduction in milk production was observed between days in milk 10 and 90 after veterinary-assisted calving compared with non-assisted calving, leading to the conclusion that non-assisted cows presented a flatter lactation curve after peak yield [69]. One of the reasons is a reduced dry matter intake in the months postpartum [86].
\nIn Jerseys the effects of calf gender in mothers milk production were not as pronounced as in Holstein-Friesians [40], which can point to a genetic selection of calf-sex biased milk production.
\nDairy calves are usually separated from their mothers right after or within hours of birth and artificially reared; therefore, the differences observed on milk production of the mother should relate to factors affecting the lactogenesis in pre- or peri-natal period [40]. The pathways through which fetal sex may influence milk production are not yet fully understood. Sex-biased milk production may reflect differential cellular capacity in the mammary gland, programmed via hormonal signals from the fetal-placental unit, or post-natal through sex-biased nursing behavior [87]. Several hypothetical mechanisms have been explored in an attempt to explain the mechanisms that may explain a sex-biased milk production in bovine, albeit with discrepant results.
\nOne possible mechanism may relate to the translocation of fetal hormones to the cow mammary gland via the maternal circulation [1]. The concentrations of sexual hormones differ between male and female fetuses and can potentially enhance or inhibit mammary milk synthesis if they get access to the maternal circulation. In the bovine species, fetal steroid hormones are present from the first trimester [1, 88, 89]. The hormones produced by the bovine fetus can cross the placenta to the cow circulation and calf sex influences hormonal levels in the mother [76, 90, 91, 92, 93, 94]. Thereby, variations in the blood levels of the hormones involved in lactogenesis may influence milk, dependent on the sex of the calf born [40]. In humans, higher concentrations of circulating androgens during the second trimester were associated with a lower probability of sustaining breastfeeding to three months post-partum, but the effect of fetal sex on the milk production was not directly analyzed [1, 95].
\nAlso, it is possible that the sex of the first parity calf affects milk production for the duration of the productive life of a cow due to the differences in the level of the hormones that influence mammary development, as it has been reported in mice [96], since dairy cows are first bred before they are fully mature, usually with only 60% of their adult weight.
\nXiang and colleagues [97] showed gender variations in the placenta weight in both Bos taurus and Bos indicus pregnancies; the placenta of the male fetus present heavier total placenta weight, better placenta efficiency heavier fetus weight than female fetus. These differences might explain and favor the fact that male calves are usually heavier than the female’s.
\nDifferences in the amount of placental lactogen produced between female and male fetus could differently prime the mammary gland of the cow [1]. It is accepted that prolactin and placental lactogens have roles in mammogenesis and lactogenesis but the mechanisms of action of those hormones act are still in discussion, and the role of the calf gender is still unclear [98, 99]. Albeit the information available for bovine is scarce, in humans, differences were found in the levels of placental lactogen in the umbilical cord blood in female and male pregnancies [100]. It was also been shown that glucose-to-insulin ratios were lower in women bearing a female vs. those bearing a male fetus [101]. Both insulin and glucose are important modulators of milk production. The fetal Insulin-like peptide 3 (INSL3) are raised in maternal circulation during pregnancy in male-pregnant dairy cows and diminished in female-pregnant cows [102]. It was also demonstrated that the level of this hormone directly affects milk production [103, 104]. In cows, Insulin and IGF-I concentrations, important metabolic mediators of the energetic metabolism and body condition, are negatively associated with milk yield during the production phase of the lactation [105].
\nHienddleder et al. [106] showed that total thyroxine concentrations were higher in male pregnancies, while triiodothyronine concentrations were unaffected by fetal gender. Contrastingly, free thyroxine concentrations were higher in female pregnancies of Bos indicus genetics, while in the Bos taurus, the values for that hormone tend to be higher in male pregnancies. No gender-associated differences were found regarding the Insulin-like growth factors in this study. The changes in the thyroid hormones’ concentrations may contribute to a different pattern in gene expression at the mammary gland, due to their galactopoietic role that sets the mammary gland´ s metabolic priority during lactation [107].
\nExploring another route, Chew et al. [108] showed that larger calves are associated with higher milk production, maybe related to higher concentrations of estrogen and placental lactogens during gestation. Indirectly, this could be one of the reasons why, in some cases, male calves are associated with higher milk production, since male calves are usually heavier at birth [109]. However, a negative correlation between birthweight and milk production during gestation was also found, leading to the hypothesis that the competition for nutrient between the fetus in gestation and the milk production for the current one would drive a diminished milk production. Yet, it cannot be ruled out that a high milk production is in itself responsible for a smaller birthweight of the calf in gestation [110].
\nWomen giving birth to daughters show upregulation of epithelial/lactocyte genes, which may be associated with increased milk yield [111]. Also, in dairy cattle a sex-biased in nitrogen and energy metabolism during the transition period was observed [112]. Higher odds exist for a male birth in cows that lose less body condition after calving [113, 114]. The depth of the Negative Energetic Balance (NEB) experienced by these cows may affect the sex-biased production of milk to favor one sex or the other. The usually higher NEB that cows go through in more intensive systems may account for the results obtained under highly intensive conditions compared to the ones obtained under less stressful management. Roche et al. [113] showed that a higher loss of body condition score by the cow was associated with a higher rate of born females. Higher milk producer cows usually lose more body condition score and have a higher rate of female calves’ gestation [114]. This might be the reason why it seems that the birth of a female is positive to milk production; however, the relationship between these factors might be the inverse, with higher producers having a higher rate of female calves [34].
\nCow’s milk production increases with the weight of the calf born [115], and male calves mean weight at birth is higher [82]. This difference in calf-sex birth weight can lead to the idea that the milk production is related to sex, when in fact it only reflects the birth weight [40]. Chew et al. [108] found no calf-sex bias in milk production when birth weight was included in the model.
\nThe sex of the calf whose birth initiates lactation can influence the milk production in the subsequent lactation because of the hormonal influences on the mammary gland development or due to the calf sex effects on pregnancy length. Also, fetal sex can influence lactation production during pregnancy because cows become pregnant at peak lactation [109].
\nIn the Cervus elaphus species, the red reindeer, dominant females give birth to a higher proportion of males than their subordinates. It is known that these dominant hinds produce higher levels of progesterone in the early days of pregnancy, and male blastocysts secrete interferon-tau earlier than females, so the hypothesis is that maternal recognition of pregnancy in dominant hinds is therefore more likely to be successful if the blastocyst is male [116]. Factors such as this at the time of maternal recognition of pregnancy in cattle could also affect calf sex, but this has not been studied yet.
\nHolstein heifers in the USA, even after administration of bST (bovine somatotropin) still produced significantly higher milk yield if they calve a female offspring, but sex-biased milk synthesis was not observed in parities two through five [1]. Even though hormones can cause sex-biased milk production, other factors such as birth, weight, lactation length and dystocia probably have a higher impact [40].
\nSexed semen produces 90% of offspring of the desired sex, but the fertility is reduced in between 75 to 80% compared with conventional frozen semen [117], because the sorting process produces a higher level of damaged to the spermatozoa [118]. Usually, sexed semen is applied more frequently in heifers, to profit from their higher fertility. Also, the heifers selected to be inseminated with sexed semen are usually the ones with higher genetic merit, so they are the ones producing the replacement animals [66].
\nThe use of sorted semen in dairy industry screws the gender ratios into the female sex, seeking the production of future genetically superior replacement animals. Under the sex-biased milk production framework, and according to some studies [1, 41, 57], it would be expected to observe an increase in milk yield in cows that calved a female in their first and eventually in the second parity. This effect would overcome any negative effects exerted by the calving of a larger male fetus (increasing the stress over nutrients partitioning between the fetus and the mother during pregnancy, and increasing the risk for dystocia) and variations in pregnancy length. On the other hand, the sorted semen being applied more often in heifers or primiparous cows, the former tending to present shorter pregnancy lengths [76], may also influence the results if the type of semen used does not enter in the model used. Attention should be paid when analyzing data from most studies, because usually the type of semen used in artificial insemination is not considered as a variable in the statistical model, which could affect the results.
\nAfter investigating the effect of sex-bias in milk production, using simulated data and considering different intensities of sexed semen in three different scenarios, two studies concluded that including sex-bias could increase profitability between €4.0 and €9.9 per cow per year [58, 119] (Table 3). On the other hand, it was also concluded that any increase in milk yield from cows calving a female calf was insufficient to warrant the use of sexed semen. The real influence of sex-biased milk production using sexed semen must be further studied before recommendations can be made into its economic impact [40]. Also, two different studies concluded that, even though there might be an effect of calf gender on a cows’ milk production, the impact was not large enough to influence profit [54] or encourage the use of sexed semen [56].
\nDifferences in milk yield per cow/year | \nWithout sex bias | \nWith sex bias | \nSimulation scenario | \n
---|---|---|---|
Milk yield (kg of ECM) | \n36 | \n48 | \nSorted semen used in 30% of heifers and 30% of cows | \n
Net return (€) | \n3.0 | \n7.0 | \n|
Milk yield (kg of ECM) | \n66 | \n99 | \nSorted semen used in 100% of heifers and 50% of cows | \n
Net return (€) | \n3.1 | \n13.0 | \n
Effects of the use of sex sorted semen on milk yield per cow/year considering two different simulation scenarios [109].
Whether or not a sex biased milk production in dairy cows exists, this bias can vary, favoring one sex or the other and, sometimes, none. It seems to favor females in intensive production systems, while in other less intensive systems this was not observed.
\nThe conflictual results obtained in different studies considering the cow may influence the sex of offspring suggest that the systems were cows are generally in good and competitive condition produce more milk for bull calves. They also seem to indicate that cows in a worse condition, or of a genetically diverging strain, apparently invest more milk in heifer calves. Up to now, conflicting reports have been presented to the scientific society, but differences among the models used make difficult to establish a clear relation between the gender of the offspring and the productivity of the cow. The different results observed are probably due to differences in the methodological approach, and the different influencer parameters used to calculate a lactation milk production, and in possible confounding factors that may not be completely identified. Also, other factors, such as different housing and feedings can have impact in calf-sex milk production bias in pathways still to be understood.
\nTo further explore this theory, additional research is needed that includes other cattle populations and correlating the investment strategy with an animal welfare index. If the calf sex effect in milk production is present in a population, selection of bull mothers and progeny tested bulls may be biased due to the offspring sex, increasing the genetic progress towards more profitable cows, if this calf-gender bias is accounted for in breeding value estimation.
\nThis work was funded by the project UIDB/CVT/00772/2020, supported by the Portuguese Science and Technology Foundation (FCT).
\nThe authors declare no conflict of interest.
"Open access contributes to scientific excellence and integrity. It opens up research results to wider analysis. It allows research results to be reused for new discoveries. And it enables the multi-disciplinary research that is needed to solve global 21st century problems. Open access connects science with society. It allows the public to engage with research. To go behind the headlines. And look at the scientific evidence. And it enables policy makers to draw on innovative solutions to societal challenges".
\n\nCarlos Moedas, the European Commissioner for Research Science and Innovation at the STM Annual Frankfurt Conference, October 2016.
",metaTitle:"About Open Access",metaDescription:"Open access contributes to scientific excellence and integrity. It opens up research results to wider analysis. It allows research results to be reused for new discoveries. And it enables the multi-disciplinary research that is needed to solve global 21st century problems. Open access connects science with society. It allows the public to engage with research. To go behind the headlines. And look at the scientific evidence. And it enables policy makers to draw on innovative solutions to societal challenges.\n\nCarlos Moedas, the European Commissioner for Research Science and Innovation at the STM Annual Frankfurt Conference, October 2016.",metaKeywords:null,canonicalURL:"about-open-access",contentRaw:'[{"type":"htmlEditorComponent","content":"The Open Access publishing movement started in the early 2000s when academic leaders from around the world participated in the formation of the Budapest Initiative. They developed recommendations for an Open Access publishing process, “which has worked for the past decade to provide the public with unrestricted, free access to scholarly research—much of which is publicly funded. Making the research publicly available to everyone—free of charge and without most copyright and licensing restrictions—will accelerate scientific research efforts and allow authors to reach a larger number of readers” (reference: http://www.budapestopenaccessinitiative.org)
\\n\\nIntechOpen’s co-founders, both scientists themselves, created the company while undertaking research in robotics at Vienna University. Their goal was to spread research freely “for scientists, by scientists’ to the rest of the world via the Open Access publishing model. The company soon became a signatory of the Budapest Initiative, which currently has more than 1000 supporting organizations worldwide, ranging from universities to funders.
\\n\\nAt IntechOpen today, we are still as committed to working with organizations and people who care about scientific discovery, to putting the academic needs of the scientific community first, and to providing an Open Access environment where scientists can maximize their contribution to scientific advancement. By opening up access to the world’s scientific research articles and book chapters, we aim to facilitate greater opportunity for collaboration, scientific discovery and progress. We subscribe wholeheartedly to the Open Access definition:
\\n\\n“By “open access” to [peer-reviewed research literature], we mean its free availability on the public internet, permitting any users to read, download, copy, distribute, print, search, or link to the full texts of these articles, crawl them for indexing, pass them as data to software, or use them for any other lawful purpose, without financial, legal, or technical barriers other than those inseparable from gaining access to the internet itself. The only constraint on reproduction and distribution, and the only role for copyright in this domain, should be to give authors control over the integrity of their work and the right to be properly acknowledged and cited” (reference: http://www.budapestopenaccessinitiative.org)
\\n\\nOAI-PMH
\\n\\nAs a firm believer in the wider dissemination of knowledge, IntechOpen supports the Open Access Initiative Protocol for Metadata Harvesting (OAI-PMH Version 2.0). Read more
\\n\\nLicense
\\n\\nBook chapters published in edited volumes are distributed under the Creative Commons Attribution 3.0 Unported License (CC BY 3.0). IntechOpen upholds a very flexible Copyright Policy. There is no copyright transfer to the publisher and Authors retain exclusive copyright to their work. All Monographs/Compacts are distributed under the Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0). Read more
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\\n\\nThe Open Access publishing model employed by IntechOpen eliminates subscription charges and pay-per-view fees, enabling readers to access research at no cost. In order to sustain operations and keep our publications freely accessible we levy an Open Access Publishing Fee for manuscripts, which helps us cover the costs of editorial work and the production of books. Read more
\\n\\nDigital Archiving Policy
\\n\\nIntechOpen is committed to ensuring the long-term preservation and the availability of all scholarly research we publish. We employ a variety of means to enable us to deliver on our commitments to the scientific community. Apart from preservation by the Croatian National Library (for publications prior to April 18, 2018) and the British Library (for publications after April 18, 2018), our entire catalogue is preserved in the CLOCKSS archive.
\\n"}]'},components:[{type:"htmlEditorComponent",content:'The Open Access publishing movement started in the early 2000s when academic leaders from around the world participated in the formation of the Budapest Initiative. They developed recommendations for an Open Access publishing process, “which has worked for the past decade to provide the public with unrestricted, free access to scholarly research—much of which is publicly funded. Making the research publicly available to everyone—free of charge and without most copyright and licensing restrictions—will accelerate scientific research efforts and allow authors to reach a larger number of readers” (reference: http://www.budapestopenaccessinitiative.org)
\n\nIntechOpen’s co-founders, both scientists themselves, created the company while undertaking research in robotics at Vienna University. Their goal was to spread research freely “for scientists, by scientists’ to the rest of the world via the Open Access publishing model. The company soon became a signatory of the Budapest Initiative, which currently has more than 1000 supporting organizations worldwide, ranging from universities to funders.
\n\nAt IntechOpen today, we are still as committed to working with organizations and people who care about scientific discovery, to putting the academic needs of the scientific community first, and to providing an Open Access environment where scientists can maximize their contribution to scientific advancement. By opening up access to the world’s scientific research articles and book chapters, we aim to facilitate greater opportunity for collaboration, scientific discovery and progress. We subscribe wholeheartedly to the Open Access definition:
\n\n“By “open access” to [peer-reviewed research literature], we mean its free availability on the public internet, permitting any users to read, download, copy, distribute, print, search, or link to the full texts of these articles, crawl them for indexing, pass them as data to software, or use them for any other lawful purpose, without financial, legal, or technical barriers other than those inseparable from gaining access to the internet itself. The only constraint on reproduction and distribution, and the only role for copyright in this domain, should be to give authors control over the integrity of their work and the right to be properly acknowledged and cited” (reference: http://www.budapestopenaccessinitiative.org)
\n\nOAI-PMH
\n\nAs a firm believer in the wider dissemination of knowledge, IntechOpen supports the Open Access Initiative Protocol for Metadata Harvesting (OAI-PMH Version 2.0). Read more
\n\nLicense
\n\nBook chapters published in edited volumes are distributed under the Creative Commons Attribution 3.0 Unported License (CC BY 3.0). IntechOpen upholds a very flexible Copyright Policy. There is no copyright transfer to the publisher and Authors retain exclusive copyright to their work. All Monographs/Compacts are distributed under the Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0). Read more
\n\nPeer Review Policies
\n\nAll scientific works are Peer Reviewed prior to publishing. Read more
\n\nOA Publishing Fees
\n\nThe Open Access publishing model employed by IntechOpen eliminates subscription charges and pay-per-view fees, enabling readers to access research at no cost. In order to sustain operations and keep our publications freely accessible we levy an Open Access Publishing Fee for manuscripts, which helps us cover the costs of editorial work and the production of books. Read more
\n\nDigital Archiving Policy
\n\nIntechOpen is committed to ensuring the long-term preservation and the availability of all scholarly research we publish. We employ a variety of means to enable us to deliver on our commitments to the scientific community. Apart from preservation by the Croatian National Library (for publications prior to April 18, 2018) and the British Library (for publications after April 18, 2018), our entire catalogue is preserved in the CLOCKSS archive.
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I am also a member of the team in charge for the supervision of Ph.D. students in the fields of development of silicon based planar waveguide sensor devices, study of inelastic electron tunnelling in planar tunnelling nanostructures for sensing applications and development of organotellurium(IV) compounds for semiconductor applications. I am a specialist in data analysis techniques and nanosurface structure. I have served as the editor for many books, been a member of the editorial board in science journals, have published many papers and hold many patents.",institutionString:null,institution:{name:"Sheffield Hallam University",country:{name:"United Kingdom"}}},{id:"54525",title:"Prof.",name:"Abdul Latif",middleName:null,surname:"Ahmad",slug:"abdul-latif-ahmad",fullName:"Abdul Latif Ahmad",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"20567",title:"Prof.",name:"Ado",middleName:null,surname:"Jorio",slug:"ado-jorio",fullName:"Ado Jorio",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Universidade Federal de Minas Gerais",country:{name:"Brazil"}}},{id:"47940",title:"Dr.",name:"Alberto",middleName:null,surname:"Mantovani",slug:"alberto-mantovani",fullName:"Alberto Mantovani",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"12392",title:"Mr.",name:"Alex",middleName:null,surname:"Lazinica",slug:"alex-lazinica",fullName:"Alex Lazinica",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/12392/images/7282_n.png",biography:"Alex Lazinica is the founder and CEO of IntechOpen. After obtaining a Master's degree in Mechanical Engineering, he continued his PhD studies in Robotics at the Vienna University of Technology. Here he worked as a robotic researcher with the university's Intelligent Manufacturing Systems Group as well as a guest researcher at various European universities, including the Swiss Federal Institute of Technology Lausanne (EPFL). During this time he published more than 20 scientific papers, gave presentations, served as a reviewer for major robotic journals and conferences and most importantly he co-founded and built the International Journal of Advanced Robotic Systems- world's first Open Access journal in the field of robotics. Starting this journal was a pivotal point in his career, since it was a pathway to founding IntechOpen - Open Access publisher focused on addressing academic researchers needs. Alex is a personification of IntechOpen key values being trusted, open and entrepreneurial. Today his focus is on defining the growth and development strategy for the company.",institutionString:null,institution:{name:"TU Wien",country:{name:"Austria"}}},{id:"19816",title:"Prof.",name:"Alexander",middleName:null,surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/19816/images/1607_n.jpg",biography:"Alexander I. Kokorin: born: 1947, Moscow; DSc., PhD; Principal Research Fellow (Research Professor) of Department of Kinetics and Catalysis, N. Semenov Institute of Chemical Physics, Russian Academy of Sciences, Moscow.\r\nArea of research interests: physical chemistry of complex-organized molecular and nanosized systems, including polymer-metal complexes; the surface of doped oxide semiconductors. He is an expert in structural, absorptive, catalytic and photocatalytic properties, in structural organization and dynamic features of ionic liquids, in magnetic interactions between paramagnetic centers. The author or co-author of 3 books, over 200 articles and reviews in scientific journals and books. He is an actual member of the International EPR/ESR Society, European Society on Quantum Solar Energy Conversion, Moscow House of Scientists, of the Board of Moscow Physical Society.",institutionString:null,institution:{name:"Semenov Institute of Chemical Physics",country:{name:"Russia"}}},{id:"62389",title:"PhD.",name:"Ali Demir",middleName:null,surname:"Sezer",slug:"ali-demir-sezer",fullName:"Ali Demir Sezer",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/62389/images/3413_n.jpg",biography:"Dr. Ali Demir Sezer has a Ph.D. from Pharmaceutical Biotechnology at the Faculty of Pharmacy, University of Marmara (Turkey). He is the member of many Pharmaceutical Associations and acts as a reviewer of scientific journals and European projects under different research areas such as: drug delivery systems, nanotechnology and pharmaceutical biotechnology. Dr. Sezer is the author of many scientific publications in peer-reviewed journals and poster communications. Focus of his research activity is drug delivery, physico-chemical characterization and biological evaluation of biopolymers micro and nanoparticles as modified drug delivery system, and colloidal drug carriers (liposomes, nanoparticles etc.).",institutionString:null,institution:{name:"Marmara University",country:{name:"Turkey"}}},{id:"61051",title:"Prof.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"100762",title:"Prof.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"St David's Medical Center",country:{name:"United States of America"}}},{id:"107416",title:"Dr.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Texas Cardiac Arrhythmia",country:{name:"United States of America"}}},{id:"64434",title:"Dr.",name:"Angkoon",middleName:null,surname:"Phinyomark",slug:"angkoon-phinyomark",fullName:"Angkoon Phinyomark",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/64434/images/2619_n.jpg",biography:"My name is Angkoon Phinyomark. I received a B.Eng. degree in Computer Engineering with First Class Honors in 2008 from Prince of Songkla University, Songkhla, Thailand, where I received a Ph.D. degree in Electrical Engineering. My research interests are primarily in the area of biomedical signal processing and classification notably EMG (electromyography signal), EOG (electrooculography signal), and EEG (electroencephalography signal), image analysis notably breast cancer analysis and optical coherence tomography, and rehabilitation engineering. I became a student member of IEEE in 2008. During October 2011-March 2012, I had worked at School of Computer Science and Electronic Engineering, University of Essex, Colchester, Essex, United Kingdom. In addition, during a B.Eng. 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