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In the recent decades, the population in the industrialized Western countries has become remarkable sedentary and have had a considerable increase in the intake of what has been called "fast food," meals that are rich in fat and carbohydrates and contain elevate levels of cholesterol as well. The elevated consumption of fast food has had a strong impact on public health, which has important repercussions in several levels including an economic impact due to the elevated cost of a chronic use of specialized health services and a detrimental effect in both, life quality and expectancy for the patients. Among the adverse health effects of this type of diet, we can mention obesity, vascular diseases, and metabolic syndrome, and it has been recently proposed that it can increase the risk of developing Alzheimer’s disease (AD), which is the most common type of dementia in elderly people. It is considered that a particular type of diet could accelerate the progress of the disease for a not yet well-known mechanism [1]. It is a revolutionary idea, since we have had for several years the conception that brain is actually protected by the blood–brain barrier (BBB); however, experimental evidence suggests that the consumption of diets rich in fat can disrupt the permeability of BBB, making it vulnerable to systemic molecules that could trigger degenerative processes [1, 2].
\nIn the current chapter, we will review the state of the art related to the impact of diets rich in fat or cholesterol on the brain, and how the alterations induced in other organs can impact brain functioning and could increase the susceptibility to develop dementia. The bibliographic revision was carried out running an exhaustive search on the research articles related to the topic employing the database of the US National Library of Medicine, National Institutes of Health, PubMed.gov. Firstly, reviewing the most recent papers and those with the most relevant information. Thereafter, we carefully followed the references cited by the reviewed articles in order to study the grounding data on the subject and which direction it followed until our days in order to document the accuracy and evolution of the data.
\nOne of the histopathological hallmarks of AD is the extracellular deposition of amyloid-β peptide (Aβ) in the brain. It is widely accepted that Aβ deposition occurs when the neuronal synthesis of the peptide exceeds the clearance capacity [3, 4]. However, some decades ago, the idea was proposed that Aβ generated systemically could pass the BBB and be deposited in the brain, since Aβ was detected in noncerebral biological fluids. Such idea raised from grounding data of Seubert et al. [5], who demonstrated that Aβ fragment comprising the amino acids 13–28 can be detected in cerebrospinal fluid and plasma of several species including human as well as in conditioned media from human brain cell cultures. It originated the idea that cerebral Aβ deposits could be generated systemically and for unknown mechanism, accumulate in the brain where they affect the capacity, to be clear, increasing the amount of the peptide and eventually form the extracellular deposits. A good amount of data has focused on this idea since then. An interesting line of study has focus on the production of Aβ by noncerebral tissue induced by consumption of diets rich in fat. One of the physiological functions of Aβ is relate to lipids metabolism and many Aβ transport proteins have been associated with lipids in vivo [6]. The association of the Aβ soluble fraction with high-density lipoproteins from healthy human plasma and cerebrospinal fluid was reported as well [7, 8]. The association between lipids and Aβ was demonstrated in a very elegant study where Aβ activity was followed labeling it with radioactivity, and it was found that the peptide is expressed in tissues rich in fat, such as spleen, marrow, liver, adipose tissue, brain, kidney, lung, and skeletal muscle. It was shown that the expression of Aβ is associated with postprandial lipoproteins such as chylomicrons, lipoproteins that are in charge to move dietary fat from intestine to the target organs. These associations remain during lipolysis and tissue uptaking processes [9]. Therefore, it can be proposed that an increased plasmatic amount of such proteins containing Aβ could produce an imbalance and could even be delivered in brain contributing to cerebral amyloidosis, one of the responsible events related to Alzheimer’s disease [9, 10]. The natural question is: how can we increase the amount of Aβ associated to postprandial lipids? One answer is the intake of diets rich in fat or cholesterol because they could break the balance of lipids content, but by which way? An interesting direction has been to study the expression of Aβ in organs rich in lipids and if such expression is regulated by fat or cholesterol diets.
\nKoudinov et al. [11] reported that hepatocytes secrete amyloid-β as a lipoprotein complex. Another organ where it has been documented that Aβ is produced is the small intestine. Given the evidence that Aβ is associated to postprandial lipoproteins, chylomicrons, Galloway et al. [10] followed this line of evidence and studied small intestinal epithelial cells (where the chylomicrons are produced). They fed wild-type mice with low- or high-fat diet. After six months of treatment they determinate by immunohistochemistry, the expression of the amyloid precursor protein in absorptive cells in the small intestine and observed a greater expression of this molecule in small intestinal epithelial cells of high-fat fed animals, whereas animals fasting 65 h did not show any expression. There is another study where the group of John CL Mamo evaluated the expression of Aβ in enterocytes after a low- or high-fat diet with 1% cholesterol in apoliprotein E (apo E) (-/-) knockout mice. Apoliprotein E is a lipoprotein that modulates Aβ biogenesis [12–14]. After six months of dietary treatment, the small intestine of apo E (-/-) KO mice fed with low-fat diet showed the same levels of expression of Aβ as the wild-type animals detected by immunohistochemistry. On the other hand, both groups of animals, wild-type and apo E (-/-) KO mice fed with high-fat diet, showed an increased expression of Aβ in enterocytes being higher in the KO animals. Also in these study, the group evaluates villi length between the groups treated, finding that the high-fat diet did not affect villi length in apo E(-/-) KO mice, but interestingly there is an increase in villi length of KO mice treated with low-fat diet when compared with wild-type mice under the same dietary conditions [15]. These groups also carried out a very elegant study to corroborate the association of Aβ production with recently generated lipoproteins, employing three-dimensional immunofluorescence microscopy and determinated that Aβ produced by enterocytes certainly has a clear colocalization with chylomicrons in small intestine enterocyte after three months of dietary treatment (free of cholesterol). They found that the amount of Aβ colocalizing with chylomicrons reaches the double [16]. These data together confirms the presence of Aβ in lipoproteins generated in small intestine and that a diet rich in fat could increase the production of transport lipoproteins. However, the open question stills remains: how this Aβ produced systemically reaches the brain? (Figure 1). Further studies are necessary to establish if indeed an imbalance in lipids production induced by diet can promote the delivery of these systemic Aβ to brain and induce cerebral amyloidosis.
\nThe ingestion of food rich on fat and cholesterol can increase the amount of postprandial lipoproteins chylomicrons. An increased production of chylomicrons can lead to an overproduction of Aβ and potentially produce an unbalance on Aβ processing and lead to cerebral amyloidosis.
The brain is a very well-protected organ with two barrier systems. One is a highly specialized microvascular endothelial system known as blood brain barrier (BBB), its function is to protect the brain from the entry of damaging substances and at the same time, allows the entry of nutrients as well as endocrine signals by means of an active transport and a passive diffusion system. The second is the choroid plexus, whose function is to prevent the entry of blood in the cerebrospinal circulation [17]. An unbalance in such systems could lead to disease conditions regarding the entrance of damaging molecules or disrupting the entrance of proper nutrients or endocrine signals. A good body of data has focused in study; how dietary habits can trigger BBB disruption? A longitudinal study, carried out in Sweden, evaluated the integrity of BBB in vivo in 81 women with a wide range of body size, who acceded to receive a lumbar puncture in order to obtain cerebrospinal fluid and compared the index of albumin content. Albumin is a constitutive protein that is absent in the brain, since its access is stopped by the BBB; therefore its presence in cerebrospinal fluid is a sign of disruption of the protection systems. Among these large group studied, the obese and overweight women between 70 and 84 years had the highest amount of albumin reported as the ratio of albumin in cerebrospinal fluid/Serum albumin (CSF/S albumin). Interestingly, they found a correlation between low levels of sex hormone binding globulin (SHBG) in the same group of women when they were younger [18]. It is known that SHBG decreases with overweight in both, male and female [19–21]. In the Swedish longitudinal study, SHBG was employed as measure of endocrine signal in the same group of females when they were in there middle forties, and decades later when they were analyzed for several parameters besides the CSF/S albumin ratio, such measures included behavioral evaluations finding that they had cognitive alterations [18]. It strongly suggests that since youth, these group of obese and overweighting women had less content of SHBG accompanied in elderly years by BBB disruption and cognitive decline. These data suggest that an unbalance between the selective entrance and exit of molecules and signaling drived by a failed BBB filtering can lead to development of dementia, but more experimental data is needed in order to elucidate the mechanism behind this effect. One way to explain the cognitive detriment found in these patients could be the diminishment of factors that have been shown to be protective for the brain, such as SHBG. High levels of SHBG have been associated with neuroprotection in stroke, vascular and cardiovascular diseases, diabetes [21–25], and an increased amount of molecules potentially damaging for the brain, such as Aβ [26–28]. Such idea can be supported by the fact that it was found in the obese and overweight women, a higher ratio of CSF/S albumin has been observed in subjects with AD as well [29, 30]. In this study, the CSF/S albumin content was measured in 118 patients diagnosed with AD and clinical data of vascular alterations was registered as well. The AD subjects were compared with individuals without dementia of the same age, finding a higher albumin ratio in those with both AD and vascular factors. There was not significant BBB disruption in the patients without vascular alterations; additionally, there is no correlation with BBB disturbances and age in the control group, which strongly suggests a relationship with the vascular alterations, BBB disruption, and AD [29]. Controversially in a study, albumin content as well as IgG in serum and cerebrospinal fluid in several groups of patients with different dementias such as early-progression familial AD, the senile dementia of Alzheimer type (Late Onset Alzherimer’s Disease LOAD), and two types of vascular dementia: a group diagnose with vascular dementia and others with multiinfarct, were measured. The multiinfarct group was reported with the highest significant alteration of the BBB but not in AD group. All these data supports the idea that vascular factors associated to BBB disruption are in relationship with the development of many dementia syndromes and are not restricted to AD [31]. That controversial information can be clarified with animal experimental data, where several variables can be controlled. The very first experimental evidence that the Aβ peptide can actually cross BBB and be deposited in the brain parenchyma was done in 1993 by Zlokovic et al. [32]. The researchers injected synthetic forms of Aβ peptide: 1–28 and 1–40, which were labeled with a radioactive marker in order to follow it after carrying out an injection in the neck vessel of the guinea pigs. The research group found a specific deposition of both synthetic peptides in the BBB microvasculature, initiating in the luminal side and transcellular transport into the brain parenchyma. This study strongly supports the idea that the Aβ produced systemically can cross the BBB. However, the mechanism remains unclear so far.
\nAlthough there is evidence that BBB can be disrupted in patients with dementia, it is possible that the development on AD can be due to systemically produced Aβ that can cross the BBB and form the deposits in the brain, but how does this happen? As we reviewed, obese patients apparently have a disrupted BBB permeability, although, what triggers that? Are the intake habits involved in such phenomenon? There is experimental evidence that suggest that components of Western diet, such as cholesterol and saturated fat, can contribute to that phenomenon. Studies with rabbits fed with a diet containing 2% cholesterol for 8 weeks, have demonstrated that such type of diet disrupts BBB permeability, alters vascularity, and induce vessels inflammation and Aβ peptide accumulation in parenchyma [26–28]; and this accumulation is similar to that observed in brains of AD patients [33]. This body of data, mainly generated by D.L. Sparks and collaborators, strongly supports the idea that high cholesterol consumption, importantly, contributes to the development of AD onset by the accumulation of Aβ, vascular alterations, as well as BBB selective permeability disruption.
\nThe contribution of BBB disruption of a high energetic diet (HE) (approximately 40% Kcal of fat versus 13% of standard laboratory rodent diet) based on high saturated fat and glucose was evaluated in 60-days-old 32 male rats that were fed for 90 days with this type of diet. The researchers evaluated the BBB integrity, measuring by ELISA, the content of sodium fluorescein (NaFl) injected throughout the femoral artery in the prefrontal cortex, striatum, and hippocampus of the treated rats. They found a significant increased amount of NaFl in the hippocampus of the treated rats compared with the control but not in prefrontal cortex or in striatum. They also measured the mRNA expression of tight junction proteins by RT-PCR in choroid plexus and BBB capillaries. Thigh junction proteins are critical components for maintenance of selective BBB permeability, its diminishment can alter the BBB function. They found a decrease expression of the thigh junction proteins and alterations in behavioral task directly associated with hippocampal function [1]. A further study was carried out by Davidson et al. [34], where they fed 24 male rats with a high energy diet as well as high saturated fat and glucose and following for different time points (7, 14, 21 and 28 days), evaluated BBB integrity by injecting NaFl following the same procedure reported by Kanoski et al. [1]. They found that the hippocampus was the brain structure that exhibit the highest concentration of the dye compared with prefrontal cortex and striatum. In this study, the researchers evaluate the differences between those animals, under HE diet, that show what they called obesity resistant versus those that developed obesity. The obesity resistant group was the one that consumed the HE diet but gained the least weight and body fat. The animals included in the obesity group were those that gained the most bodyweight and fat. It was this last group that showed the major BBB permeability and had the highest deposit of NaFl in the hippocampus. Interestingly, they found that those animals, in the HE diet, had the lowest bodyweight and the lowest amount of fat, and did not show difference in the behavioral performance compared with the control group. However, those rats that developed obesity and had the higher deposit of dye in the hippocampus, showed alterations in the performance of the hippocampal-dependent tasks [34]. These evidences directly shows a relationship between diets rich in fat, obesity development, and hippocampal-related cognitive alterations. We will discuss in the next section, the relevance of the hippocampal structure, cognitive performance, and its detriment.
\nFrom the information reviewed in this section, we can conclude that BBB alteration is a feature that takes part of dementia onset in both, AD and vascular dementia. Obesity can contribute to this phenomenon and, although the mechanism is not well known, a particular factor that can participate in this process is the intake of diets rich in cholesterol or fat, as well as glucose, those known components of a typical Western diet (Figure 2).
\nThe overproduction of systemic Aβ, produced by consume diets rich on fat or cholesterol, can promote and alter the selective permeability of BBB, allowing the passage of molecules to the brain, such as systemic Aβ that was not clear and lead to cerebral amyloidosis and brain inflammation.
The hippocampus is a brain structure considered as a part of the allocortex, which is one of the oldest brain areas from the phylogenetical point of view. It has a high capacity of plasticity; it is directly involved in learning and memory process and, interestingly, is very susceptible to damage and has attracted the research focus for several years since it is one of the first brain structures that degenerate during the AD process [35]. As we reviewed in the last section, the hippocampus seems to be very susceptible to the effect of consumed diets rich in fat or cholesterol, but can this actually drive the brain into a degenerative process? Can it contribute to the development of dementia? We will discuss this idea in the current section. First, we will review how the diet high in cholesterol or fat can contribute to the development of features associated with AD, particularly with amyloidosis.
\nDiets rich in cholesterol, as we have reviewed, can induce vascular inflammation, BBB, and promotes Aβ peptide accumulation in the brain parenchyma in an animal model of rabbit fed with high-cholesterol diet [26–28]. Supporting the association of elevated concentrations of cholesterol and AD detriment in a very recent in vitro study carried out by Avila-Muñoz and Arias [36] in isolated astrocytes obtained from brain cortex of 1- to 3-day-old Wistar rats, they found astrocyte activation, An increase on the expression of amyloid precursor protein (APP), and promoted its amyloidogenic processing, and an increase in reactive species oxygen (ROS), a marker of oxidative stress, after treating the culture for 48 h with cholesterol concentrated at 25 or 50 μM. All these parameters measured, including glia activation, resemble features that have been found in postmortem brain tissues obtained from AD patients [37–39], but how the consumption of a diet high in cholesterol can contribute to the development of AD? In vivo studies can answer this question. Transgenic mice Tg2576 (which express the human APP695 carrying the Swedish double mutation at codons 595 and 596, Hsiao et al. [40]), were fed with a 5% cholesterol diet for 6 weeks. They found an increase of the APP cytosolic fragment but apparently the hypocholesteremia induced by the diet does not deregulates Aβ metabolism (George et al, 2004).
\nIn a further work, carried out by Refolo et al [41], with 5-months-old double-mutant for presenilin (PS) and amyloid precursor protein (PSAPP) mice, which express familial mutant PS1M146V and the APP695 mutations [42], evaluated the effect of a combined diet with 5% cholesterol and 10% fat for 7 weeks. They found that the dietary treatment induced elevated levels of cholesterol in both, plasma and brain, which is an important data since it showed that brain cholesterol is produced in situ, and this data demonstrates that brain cholesterol is increased by diet. This increase in brain cholesterol correlates with an increase of total Aβ in brain. In addition, there was an enhanced amount of Aβ, particularly not in Aβ 1–40 and 1–42, but in 1–30 and 1–34 as well. This was accompanied with an increase in the number of Aβ deposits as well as an increase in the plaque area in the hypercholesteremic transgenic mice. Interestingly, there were no changes found in presenilin 1 (PS1) processing. These data strongly supported the hypothesis that a diet high in fat and cholesterol can contribute to the development of amyloidosis, one of the main conditions to develop AD.
\nAs result of consume diets high on fat and cholesterol there is an increase levels of brain cholesterol and systemic cholesterol. Also elevates Aβ production in brain and its deposit and increases as well the glia activation and production of ROS in brain. All these together can lead to AD onset.
All these data shows experimental evidence linking the consumption of diets rich in fat and/or cholesterol with the development of amyloidosis. Nevertheless, dementia is a more complex syndrome, comprised of many other features such as cognitive decline and neuronal lost. Particularly in the hippocampus, which is as we mentioned before, one of the first areas affected during the neurodegenerative process, its susceptibility to suffer alterations resulted from consuming diets high in fat or cholesterol appears crucial as one of the possible mechanism involved in the development of AD (Figure 3). We will discuss that idea in the section below.
\nIn the last sections, we have discussed how the consumption of diets rich in fat or cholesterol can contribute to the production of Aβ peptide in noncerebral tissue. The impact that this could have in the BBB selective permeability and its participation in brain amyloidosis conditions that can contribute to the dementia onset but, besides these alterations, one of the main conditions found in dementia patients is brain atrophy and behavioral alterations. Is brain functionality affected by the components typically found in the Western diet? Could diet composition affect brain architecture and plasticity? Moreover, is cognition affected the consumption of diets rich in fat or cholesterol? We will review such ideas in the current section.
\nA link between cognitive decline and dietary habits has been proposed. There is an epidemiological study carried out with Japanese men living in Hawaii compared with age-matched men living in Japan that evaluated the prevalence of dementia employing the Diagnostic and Statistical Manual of Mental Disorders. The results found that those subjects living in the USA have a higher prevalence of dementia: 9.3% for all type dementia, 5.4% for Alzheimer’s disease, and 4.2% for vascular [43]. Continuing in this line of evidence, there is another study that was carried out with people from same ethnic background living in their natal land or in a foreign country (USA). They found in concordance with the study cited before, that those individual living in Indiana (where the study was carried out on) had a higher prevalence of dementia compared with age-matched individuals living in Nigeria or Ibadan [44]. This data strongly suggests that there are some stimuli in this Western country, which contribute to the development of several types of dementia, and the question is: what are these stimuli? A good candidate are the nutritional habits. In Western countries, especially countries such as the USA or Mexico, people consume food with high amounts of saturated fat and cholesterol and show the highest rates of obesity worldwide. The brain is an organ rich in lipids and essential fatty acids that are mainly obtained from food and have a crucial participation on brain functioning [45]. So, to think that lipids elevation induced by diet could be in detriment of the brain, which is a logical assumption, but what are the cellular mechanisms involved in the possible detrimental effect of food components such as fat and cholesterol? Well, experimental work has demonstrated evidence of the interplay between obesity, brain alteration, and cognitive decline, more especially with hippocampal-related cognitive processes. Seminal works in this area were carried out in the University of Toronto by Greenwood and Winocur [46]. They fed 1-month-old Long-Evans rats with two types of high-fat diets containing 40% of calories: a saturated fatty acids (lard-based) or a polyunsaturated fatty acids (soybean oil-based) and compared with a standard laboratory diet containing 4.5% of fat. They tested learning and memory abilities in the rats after 3 months of dietary treatment with the radial arm maze test, the variable-interval delayed alternation task, and the Hebb-Williams maze series. These tests evaluate spatial learning and memory performance and report failures in working or reference memories. They found that those animals fed with the lard-based diet showed impairment in all the tests. Following this line of evidence, they analyzed further with different types of saturated fatty acid diets: monounsaturated (MUFA) and polyunsaturated (PUFA) fatty acids, finding a direct relationship between the 3 months consumption of saturated fatty acids and failures in basic alternation rule, and remembering trial-specific information over time in the variable-interval delayed alternation task. Interestingly, they found alterations in brain’s phosphatidylcholine fatty acid profile. However, the changes in the membrane did not correlate with cognitive alterations [47]. It suggested there is another mechanism elucidating the cognitive impairment related to consumption of diets rich in fat; a good candidate is brain inflammation. Chronic inflammation is one of the principal altered events associated with AD [48], and it has been linked to obesity and has been reported that there is a correlation between both, obesity and AD [49, 50]. Middle-aged C57BL6 male mice were fed for 21 weeks with chow equivalent to Western diet containing 41% fat or a high-fat lard diet containing 60% fat for 16 weeks. They showed an alteration on learning acquisition measured by the Stone T-maze and it is accompanied by microglial activation, increase expression of cytokines like TNFα, IL-6, and MCP-1, and a decrease on brain-derivated neurotrophic factor (BNDF) [51]. Interestingly, there was not any detrimental effect observed in those animals that consumed the like-Western diet. These data agree with results from Greenwood and Winocur and propose a possible way underlying the effect diet, which is an inflammation process, and the decrease on neural factors crucial for learning processes. The results demonstrate that diet can interfere with learning abilities, but is it everything behind the diet effect on cognitive decline? There is a report with 344 white middle-aged male Fischer rats. The researchers evaluate the effect of a diet high in cholesterol and fat (diet containing 2% cholesterol and 10% hydrogenated coconut oil). The results showed a failure in working memory, here evaluated with the water radial arm maze as well as elevated lipids profile and reduce expression of Map-2 as an indicator of alteration of dendritic integrity, which correlates with memory mistakes measured in the test, and increase in inflammation markers such as microglia activation [52]. In a study carried out with Sprague-Dawley rats, which were fed for 7 days with high fat and fructose, several hippocampal alterations, such as decreased insulin signaling, were reported. In addition, they found that treated animals had a decrease in hippocampus total weight in addition with some other morphological alterations such as a diminishment on the number of dendritic spines and a reduction in the complexity of the hippocampal dendritic arborization. Moreover, there was a decrease in the expression of the microtubule-associated protein 2 (MAP-2) and in the content of synaptophysin in the CA1 region concomitant with an increased phosphorylation of tau protein, and in the presence of reactive astrocyte associated [53]. It directly demonstrates alterations in hippocampal cytoarchitecture that definitively have a strong impact on brain functionality, especially in hippocampal-related learning and memory processes.
\nAnother feature which affected by consuming diets rich in fat is adult hippocampal neurogenesis (AHN). Adult neurogenesis is a highly specialized plasticity phenomenon that, under basal conditions, occurs in two restricted brain areas: a) the subventricular zone and b) the hippocampal dentate gyrus [54, 55]. Hippocampus is a crucial area for memory processes, since its decrease is associated to memory failures, especially in short-term memory, spatial memory, and learning flexibility [56–59]. The AHN is a complex process that comprises several developmental steps starting from the division of an endogenous neural precursor cell followed by its expansion, differentiation, and fully integration to the hippocampal network [60]. These steps are reported as number of proliferative cells measured by markers of cell division; cell fate decision with the marker of early differentiation, the cytoeskeleton protein doublecortin (DCX) that is expressed in newly differentiated cells, and with NeuN, a nuclear marker of granular cells when the cell is fully differentiated. It has been recently documented that there are some food components which can regulate the neurogenic process (for a review [61]). The hippocampal neurogenesis has captured the attention since it was described in 1965 by Dass and Altman [62] due, as we already mentioned, the hippocampus is closely related to memory as well as neurodegenerative processes. Juvenile male and female Sprague-Dawley rats under a dietary regimen of high- (42% coconut butter and corn oil fat) or low-fat diet (10% fat by energy) or standard laboratory chow for 4 weeks, was found that males under high-fat diet show less cell proliferation than females and reported elevated levels of corticosterone, a stress hormone [63]. Differences in AHN were studied in mice susceptible to develop obesity (C57BL/6N) and obesity resistant (C3H/HeN). They were fed with high- and low-fat diet finding that those animals that developed obesity and consumed the high-fat diet had much lower number of proliferative cells and cells committed to neural linage (DCX positive cells), which establish a clear link between obesity and AHN diminishment [64]. In our laboratory, we have observed that 8 weeks of diet rich in fat (60%) or high in cholesterol (1.4%) in 5-months-old male Wistar rats has an impact on AHN in both, cell proliferation and more especially in the morphology of DCX cells. These cell populations have less processes and a poor complexity than animals under normal laboratory diet, and we found alterations in short-term memory (Leal-Galicia and Meraz-Ríos data not yet published). All these data together strongly suggest a detrimental effect on diets rich in fat or cholesterol in cognitive components such as navigation memory, working memory, acquisition learning, and short-term memory suggesting as mediators, alterations in brain cytoarchitecture and AHN, and associates obesity with such cognitive alterations strongly supporting the hypothesis that obesity can lead to development of dementia (Figure 4).
\nThe intake of food with high amounts of fat or cholesterol produces alteration in the hippocampus such as: reduced expression of Map-2, reduction on the number of dendritic spines and in the complexity of the dendritic tree and a decrease on neurogenesis. Consume diets with these components has also a functional impact in short-term memory, working memory and learning flexibility, that could contribute to the detriment observed in the dementia syndrome.
The consumption of diets rich in cholesterol, fat, as well as another components (carbohydrates) of the so called “Western diet” can contribute to increase the production of the peptide Aβ. This could contribute to brain amyloidosis by means of alteration of the selective permeability of the BBB, since BBB alterations are induced for these type of diets. In addition, it has been shown in the brain of transgenic animals that the amyloidosis can be accelerated by the intake of fat or cholesterol, which can lead to accumulation of Aβ in the brain. Besides that, the intake of fat or cholesterol can induce alterations in brain morphology and plasticity accompanied by a detrimental in cognitive abilities in animal models that resemble those alterations in cognitive abilities reported in AD patients, such as short-term memory, working memory, and learning flexibility. These evidences strongly suggest an association with the dietary habits and the possible development of AD in both cases, Early Onset Alzheimer’s Disease or Late Onset Alzheimer’s Disease, and a connection with systemic disruptions and brain functions (Figure 5).
\nDiets rich on fat or cholesterol that are widely consume in Western countries can lead to develop dementia onset for several ways. One is the overproduction of systemic Aβ that can reach the brain due the chronic consume of these food components can affect the selective permeability of BBB. It can facilitate the pass of systemic Aβ as well as another molecules producing brain inflammation and Aβ deposits. It is accompanied for alterations in hippocampal plasticity and its cytoarchitecture. That can have an impact on brain functionality observed as memory failures. All these together can contribute to the development of dementia.
Rice is such an agricultural commodity that covers the third-highest worldwide production making it one of the most important cereal crops [1]. With its wide geographic distribution extending from 50°N to 35°S, rice is expected to be the most vulnerable cultivated crop to changing climates in future [2, 3]. Rice production is dwindled mainly because of biotic and abiotic stresses due to the complexity of interaction between the stress factors and various molecular, biochemical and physiological phenomena affecting plant growth and development [4, 5]. To battle with these situations, development of adaptive rice varieties is one of the best strategies. Since aboveground parts are often taken into consideration for making stress tolerant varieties, root study remains backward in this aspect. Roots, the hidden portion of the plant have not yet been much focused. Because exploring the root traits of the plant are much more difficult compared to its above-ground traits. But when it comes to the fact of studying the optimal developmental plasticity system and characteristic features of plant growth, the root system is given the first priority [6]. Root system is the site of water and nutrient uptake from the soil, a sensor of abiotic and biotic stresses, and a structural anchor to support the shoot. The root system communicates with the shoot, and the shoot in turn sends signals to the roots [7]. Soil type, moisture and nutrients all strongly influence the architecture of the root system [8, 9, 10]. Recently it has been emphasized that root architectural traits play a decent role for the adaptation of crop varieties under different abiotic stresses [11, 12]. Root interaction with changing environment is a complex phenomenon that differs with genotypes and intensity of stress [13, 14, 15, 16, 17]. For that, different species and also genotypes under the same species may respond contrarily under stress conditions and show different magnitudes of tolerance or susceptibility to stress. These diversities can be exploited by plant breeders to improve stress tolerance in plants. Scientists assume that selection for yield will indirectly select for varieties with the optimum root system. But the fact is, more directed selection for specific root architectural traits could enhance yields for different soil environments [18]. As by 2035, a predicted 26% increase in rice production will be essential to feed the rising population [19], it is imperative to develop high yielding rice cultivars with efficient root systems for better exploitation of natural resources under stressed conditions.
\nBeing the hidden half of the plants, the root system performs several functions like water and nutrient acquisition, mechanical support to the plant and storage of reserve assimilates [7]. In plant, roots are the first organ for sensing the water limitation and the roots are also the signal transmitter to other plant parts through xylem sap and phytohormone which is known as one of the most important root-shoot stress signal mechanism [20, 21, 22, 23]. Development of the root system is a major agronomic trait and proper architecture in a given environment permits plants to survive in water and nutrient deficit conditions and gives the ability to utilize minimum resources efficiently [6].
\nCrop loss in rice production has become severe now-a-days due to abiotic stresses. Therefore, having a clear knowledge about the architecture and development of roots of rice toward optimizing water and nutrient uptake has become crucial for exploitation and manipulation of root characteristics for enhancing yield under unfavorable conditions [24, 25]. In general, root study comprises the study of the entire root system or a large portion of the plant’s root system [26, 27]. To understand the functional characteristic of root system and the necessity to exploit heterogeneous environment, root architecture study has become crucial in plant productivity as root system architecture is strongly linked with plasticity to the plant through which plant can alter its root structure according to its heterogeneous environment [26].
\nElongation and branching are the mode of plant root growth. Local environmental conditions, physiological status of the plants and the type of root determine the magnitude and direction of root elongation [6]. Root system architecture (RSA) is thus the three-dimensional geometry of the root system including the primary root, branch roots, and root hairs [6, 26, 28, 29]. Topological, distributional and morphological features combine to form the root system architecture [8, 26, 30]. Topology denotes the branching pattern of individual roots including features like lengths and diameters, number of roots originating from a node, root insertion angles, magnitude and the altitude of root [31, 32]. Measures of the spatial distribution of roots simplify the dissection of root systems [26]. Root morphology refers to the external features of a root axis and may include properties of roots hairs, root diameter and trend of secondary root emergence. Acceleration or inhibition of primary root growth, increment of lateral roots (LRs) and a rise in root hairs and also the formation of adventitious roots are the ways of modification of root system architecture. The primary root is formed during embryogenesis. This primary root produces secondary roots those in turn produce tertiary roots [6, 33]. Root system architecture has proved to be a critical factor in plant survival, contributing to water and nutrient acquisition efficiency and competitive fitness in a given environment [34]. Composition of soil specially water and mineral nutrients availability and plant species have impact on root architecture [6].
\nMonocot cereals have a complex fibrous root system consisting of an adventitious root (ARs) bunch. Adventitious roots originate from the shoot or subterranean stem. This type of root is sometimes referred to as a nodal or crown root [35]. Root systems of rice plants (Oryza sativa L.) comprise numerous nodal roots of relatively short length: a mature rice plant usually has several hundreds of nodal roots, most of which are less than 40 cm in length [36]. Rice (Oryza sativa L.) is a model cereal crop with seminal roots that die during the growing period [36]. Thus, lateral roots and adventitious roots are the key determinants of nutrient and water use efficiency in rice [37].
\nSeveral embryonic and postembryonic roots including the radicle, the embryonic crown roots, the postembryonic crown roots, the large lateral roots (L-type), and the small lateral roots (S-type) [38] form the rice root systems (see Figure 1). Lateral rice roots can appear on any primary root, including embryonic and crown roots, and can be classified into two main anatomical types [39]. Numerous small lateral roots (S-type) are thin with determinate growth that can be formed from large lateral roots (L-type) and they never bear any lateral roots. Whereas large lateral (L-type) roots are few in number, thinner compared to primary roots that show indeterminate growth. Additionally, lateral elongation of small lateral roots and downward elongation of large lateral roots indicate non-responsiveness of the small lateral roots to gravity. Higher orders of branching can also be observed in the large lateral roots of the crown roots that emerge at later growth stages [40]. These small and large lateral roots exhibit differential growth and lateral root bearing pattern signifying unlike purposes for these two types of lateral roots [37].
\nA typical root system architecture at the tiller axis of Oryza sativa L. Black disks indicate individual root bearing phytomer with progressive development chronologically from top to downward. Root hairs form on main axis and all the lateral roots [41].
The concept of a phytomer was established around 6–7 decades ago [40, 42]. Clear knowledge about phytomer is required for better understanding of plant development and architecture. Many higher plants, including rice, are composed of successive stem segments called phytomer [43, 44, 45]. Each phytomer consists of an internode of the stem with one leaf, one tiller bud and several adventitious (nodal) roots [36]. The phytomer concept has long been recognized among grass scientists [46, 47]. The coordinated development of stem, tiller bud, and adventitious roots in each phytomer corresponds to the phyllochronic time in rice [43, 44, 48]. This indicates that genotypic variation in root-and-shoot growth can be ascribed to the variation of stem and adventitious root development at the phytomer level [49].
\nDetailed study of root morphology and architecture at the phytomer level become more obvious with the attainment of new knowledge about segmental architecture of poaceous crops [50, 51, 52, 53]. As the higher plant structure is formed by the repetitive unit of plant growth called phytomer [54], so phytomer formation, its growth and senescence ultimately determine development of plant canopy [47]. Therefore the phytomer components have become the interest of the plant breeder.
\nRoot axes of rice plants serve functions of anchorage and typically establish overall root system architecture [55]. The lateral roots are the functionally active part of the root system involved in nutrient acquisition and water uptake. The size, type and distribution of lateral roots eventually decide the ultimate length and surface area of an individual root and finally of a whole tiller. Understanding morphology of the lateral roots is therefore important to develop rice cultivars with an efficient root system [11, 56].
\nIn rice, there are two types of lateral roots; long and thick roots, and short and slender roots [57, 58, 59]. It has been designated that the first type as L-type and the latter as S-type [60]. The L-type lateral roots are usually long and thick and are capable of producing higher-order lateral roots, whereas S-type ones are short, slender, and non-branching. In rice plants, these two types of lateral roots are visually distinguishable. The L-type lateral roots show basically identical tissue arrangement with seminal and nodal roots, whereas S-types are anatomically different wherein their vascular systems are simplified [35].
\nIn rice plants, the observed average diameter of S-type lateral roots (first-order) that were produced on mature nodal roots of a one-month-old plant was 80 μm, whereas that of L-type roots was almost double that, i.e., 159 μm. Average length was 7.6 mm for S-type and about 30 mm for L-type. The S-type laterals were almost similar in length, and only very few S-type laterals exceeded 10 mm in length. The L-type laterals varied greatly in length and some of them elongated to more than 300 mm [60]. The small laterals are less effective in water and nutrient uptake than even root hairs [61].
\nThe changes in lateral root development were triggered by changes in water status in the root zone, and these developmental changes were induced by genetic [62, 63] and environmental factors. With regard to the environmental factors, it is shown that phenotypic plasticity promoted lateral root development and that nodal root production was the key trait that ensured stable growth of rice plants grown under changing soil moisture levels [64]. As far as the literature explored, developmental morphology of the individual roots with special reference to different lateral root branches was not studied in detail, probably due to lack of the most appropriate tools and methods [11].
\nRoot hairs are tubular-shaped cells that arise from root epidermal cells called trichoblast; they are thought to increase the absorptive capacity of the root by increasing the surface area [65]. Root hairs contribute as much as 77% of the root surface area of the cultivated crops, forming the major point of contact between the plant and the rhizosphere. Root hair is a long and narrow tube like structure originating from a single cell through tip growth (the deposition of new membrane and cell wall material at a growing tip). For being the major water and nutrient uptake site of plants, root hairs form a progressively significant model system for development studies and cell biology of higher plants [66]. Root hairs had the highest contribution toward total length and surface area of an individual root whereas main axis and first order laterals mostly contributed root volume [11].
\nRoot hairs are localized for many water channels [67], phosphate [68], nitrogen [69], potassium [70], calcium [70], and sulfate transporters [71], all of which are beneficial to water and nutrient uptake by plants [72]. There is significant inter- and intra-specific variation exists for root hair traits, and this has been linked to P uptake. Plants with longer, denser root hairs exhibit greater P uptake and plant growth in P-deficient soils [73, 74, 75]. So, the root hair traits, especially root hair length can be exploited in breeding for improved nutrient uptake and increased fertilizer use efficiency [76]. Considerable researches support an important role for root hairs in P attainment [73, 74, 75, 77, 78]. Root hair length and root hair density (which is usually correlated with root hair length) have clear value for the acquisition of P and probably other diffusion-limited nutrients such as K and ammonium [79].
\nUsually root hair traits have a low heritability and their expression is influenced by soil type resulting in lack of research in this field [6, 80, 81]. It has been proposed that plasticity in root epidermis development as a response to a variety of environmental conditions might reflect a function of root hairs in sensing environmental signals, after which plants adjust themselves to the stress conditions, such as by increasing nutrient acquisition and water uptake or by helping to anchor the plant to the soil [82, 83, 84, 85, 86, 87]. Root hair elongation increases root surface area. Root surface area increment is a common phenomenon when the plants are subjected to the stress condition like salinity, drought or other abiotic stresses [79, 88, 89, 90, 91].
\nPlants recurrently face several stresses like salinity, drought, submergence, low temperature, heat, oxidative stress and heavy metal toxicity while exposed to the nature. Growth and grain production in cereals is often limited by these stresses under field conditions. All these stresses either directly or indirectly impose osmotic stress to plants that ultimately affect the final yield of rice. Root is the first part which can sense these stresses better than other plant parts. So researchers prioritize the fact of understanding the root adaptive responses of plants upon osmotic stress. In the last 30 years, comprehensive studies have been performed focusing on architecture and developmental morphology of roots and their genetic and molecular basis [11]. Morphological and anatomical development of the rice root system was thoroughly reviewed [92] whereas the mystery of root length was also reviewed [93]. A recent study highlighting the growth, development and genetic reasons of root morphology and function of crop plants was provided by [94]. An outstanding study on root system architecture and its molecular and genetic background also greatly contributed to the relevant literature recently [37]. The physiological background of root branching was also studied [7, 33]. The root parameters that are focused by the studies comprising root anatomy, plant height, root-shoot ratio, length, diameter, density, surface area and volume of root, root elongation rate, root branching, expansion of root regarding tiller development, maximum root depth, distribution pattern of root in soil column, root hydraulic conductivity, hardpan penetrability, all of which possess innumerable functional implication [95]. Roots of large diameter show greater penetration ability [96, 97, 98] and branching [8, 99] because of having larger radii of xylem vessel and poorer axial resistance to water flux [100].
\nWater is essential for survival and plant growth. As a sessile organism, plants constantly encounter water deficit, which is the most severe environmental stress limiting plant growth and productivity in natural and agricultural systems [101, 102]. Thus, water stress tolerance has been a fundamental scientific question in plant biology.
\nPlants have evolved complex adaptive mechanisms that enable them to survive drought conditions. Over more than five decades, researchers have identified osmotic adjustment, antioxidant protection, and stomatal movement as key adaptive mechanisms for survival where both osmotic adjustment and reactive oxygen species (ROS) are involved in this plastic development process [103]. To cope with the changing water status in the growing environment, plants have evolved various adaptive mechanisms by which plants can modify root allocation and root system architecture to obtain more water [104].
\nNumerous studies have provided evidence to show that when plants are subjected to water stress, root growth is strongly inhibited, although root development is less sensitive to water stress than that of shoots [105, 106, 107].
\nRoot system architecture is regulated by osmotica [108]. The osmotic potential of the soil alters the depth of the root system, its overall mass, the rate of root elongation and the number of lateral roots in many plants, including Arabidopsis [8, 9, 107, 109, 110].
\nRoot length, root dry weight, and root production are limited by drought stress [111, 112]. Roots are the significant plant part which increase plant adaptability power to soil water deficits by maintaining water uptake under dry conditions [113]. Root and other root components such as root hair, root-shoot ratio, and root length are found to be decreased in drought sensitive varieties. But the resistant varieties which possess tolerance capacity against drought showed increase in root hair, high root to shoot ratio and root length [114]. Roots are considered as the most efficient plant organ which helps plant to uptake water and minerals from the soil and during drought stress. Root proliferation and changes in root parts occurs to take more water from deeper regions of the soil [25]. Different types of changes are observed in root growth of drought resistant rice varieties such as a deeper and highly branched root system than drought- sensitive varieties [115]. Plant also extends its roots for more nutrients (such as phosphorus) and water uptake which results in more root to shoot ratio [116]. In recent years breeding for developing larger and more efficient root systems has become the hotspot in research in some crops such as rice, as there is a relation between root system size and tolerance to water stress [81, 117].
\nThe change in lateral root development, i.e. in the plasticity of the root system, exhibited under water deficit conditions may play an important role in drought stress tolerance [35]. From an agronomical view, the knowledge about lateral root development is useful for breeding varieties with drought stress tolerance [118].
\nThe importance of root system structure is particularly recognizable when its significance in relation to its function is clearly identified. The significance of root system structure in nutrient and water uptake was stressed in previous study [119].
\nUnder waterlogged conditions, the plant roots have to function in anaerobic soil, and there are at least two morphological adaptations that roots exhibit in response to anaerobiosis, i.e., development of new adventitious roots [120, 121] and superficial rooting (i.e., the concentration of new root growth in the upper layers of the soil) [122]. Nodal root production (increase in number) continued to take place, however, in the sense that when adventitious roots in the lower nodal position of the plant’s stem die due to waterlogging injury, new adventitious roots appear at the next highest nodal position. There appears to be a direct relationship between the death of older adventitious roots and the development of new ones. Progressively waterlogged plants generally show smaller root system size than those grown in a well-drained condition. It is considered that the turgor pressure affects the cell elongation and growth of plants [123, 124]. Aerobic cultivars of rice have greater ability for plastic lateral root production than irrigated lowland cultivars under transient moisture stresses [125].
\nWe have a little understanding of the responses of roots and root hairs to salinity stress and their function in stress tolerance. The efficient root system can either avoid or lessen the osmotic stress. Usually, growth, morphology, and physiology of the roots alter first under salinity stress and the whole plant is then affected. Therefore, the responses and characteristics of the roots under saline conditions are of primary importance for plant salt-tolerance [126]. It is supposed that root morphology affects salt accumulation around the roots impeding uptake of water from saline areas. Modification of root morphology has a big potential to develop crop salt tolerance [127]. Root hairs have higher sensitivity to salt than other root traits and shoots [128]. Environmental factors also regulate the root hair development [128]. The development of root epidermal cells has great plasticity where the differentiation programs can be switched from one to another in response to external factors [17]. Plasticity in development of root epidermis as a response to a variety of environmental conditions might reflect a function of root hairs in sensing environmental signals, after which plants adjust themselves to the stress conditions [82, 84, 85, 86, 87, 129].
\nRoot hair growth and development and their physiological role in response to salt stress are largely unknown [128]. The development of root epidermis cells has great plasticity where the differentiation programs can be switched from one to another in response to external factors [17]. Root hairs have higher sensitivity to salinity than do roots and shoots [128]. Systematic study on root hair plasticity induced by salt stress and the possible role in plant adaptation/tolerance to salinity is still lacking [128]. Usually root hair traits have a low heritability and their expression is influenced by soil type resulting in lack of research in this field [6, 80, 81].
\nEarlier many scientists had reported root morphology and its distribution were greatly varied based on genotypes of plant species [13, 14, 15, 16]. There is widespread evidence that root architecture and different root characteristics of many crop species varies among genotypes [14, 130, 131, 132, 133]. In a few quite recent studies, the importance of studying root architectural traits has been emphasized for the adaptation of the crop varieties to various abiotic stress conditions. Genotypic variation has a significant role in adapting the adverse environmental and edaphic effects [14]. Inter- and intra-species variations in root architectural traits are very useful to breed the crops for root features optimum for diverse environmental conditions [134, 135, 136].
\nRoot anatomical and morphological traits have been well studied in rice [92]. Varietal differences in root morphological characteristics such as length and thickness have been reported in cultivated rice (Oryza sativa L.) in various studies [11, 14, 41, 137]. In general, the roots of upland rice cultivars are thicker and penetrate more deeply into the soil than those of lowland cultivars [14]. Root distribution has also been quantitatively characterized by using several traits, including root length, volume, and density in the soil at different depths, and these characteristics differed among cultivars [92, 138, 139, 140].
\nUnderstanding and improvement of root system and its genetics plays a pivotal role to become self-sufficient and to achieve sustainability in rice production. Actually more yields from the limited input rely on our capability to unambiguously manipulate the plants. And exploring the diversity of root architecture both in genetic and phenotypic basis will directly connect to this concern. Although great strides have been made to understand the root morphology but in future, more intense investigations to elucidate the functional implication of root morphological variation may aid in selection of root system with anticipated characteristics.
\nFuture exploration of stress responses regulated by roots at cellular or tissue level will open the door of further breeding research. Besides the modern gene pools, exploration of genes and alleles in wild relatives and landraces will also provide interesting features that will be easier to transfer to cultivated rice. Further it is important to have a better understanding on the epigenetic regulation of roots and root development under stressful conditions. There will be a need for high throughput phenotyping systems coupled with automated data analysis for accelerating the development. Endorsement of approaches including both root ideotype-based screening and selection for grain yield may establish a fruitful screening system. Alongside designing new genetic screening methods based on a better knowledge of the integrated stress responses will be also appreciated. Dynamic root/soil interaction modeling will aid in integrating different functional parameters (e.g. water uptake per length of root) under a variety of environmental conditions. Overall the root system being less accessible and more complex than other agronomic traits, achieving the ambitious goal of future rice root research, coordinated effort and joint resources are required. The sensible and appropriate efforts will have a crucial role to play in future crop production in vulnerable climate and resource scarcity prioritizing the objective of serving food to 9 billion world populations by the year 2050.
\n“The authors declare no conflict of interest.”
Edited by Jan Oxholm Gordeladze, ISBN 978-953-51-3020-8, Print ISBN 978-953-51-3019-2, 336 pages,
\nPublisher: IntechOpen
\nChapters published March 22, 2017 under CC BY 3.0 license
\nDOI: 10.5772/61430
\nEdited Volume
This book serves as a comprehensive survey of the impact of vitamin K2 on cellular functions and organ systems, indicating that vitamin K2 plays an important role in the differentiation/preservation of various cell phenotypes and as a stimulator and/or mediator of interorgan cross talk. Vitamin K2 binds to the transcription factor SXR/PXR, thus acting like a hormone (very much in the same manner as vitamin A and vitamin D). Therefore, vitamin K2 affects a multitude of organ systems, and it is reckoned to be one positive factor in bringing about "longevity" to the human body, e.g., supporting the functions/health of different organ systems, as well as correcting the functioning or even "curing" ailments striking several organs in our body.
\\n\\nChapter 1 Introductory Chapter: Vitamin K2 by Jan Oxholm Gordeladze
\\n\\nChapter 2 Vitamin K, SXR, and GGCX by Kotaro Azuma and Satoshi Inoue
\\n\\nChapter 3 Vitamin K2 Rich Food Products by Muhammad Yasin, Masood Sadiq Butt and Aurang Zeb
\\n\\nChapter 4 Menaquinones, Bacteria, and Foods: Vitamin K2 in the Diet by Barbara Walther and Magali Chollet
\\n\\nChapter 5 The Impact of Vitamin K2 on Energy Metabolism by Mona Møller, Serena Tonstad, Tone Bathen and Jan Oxholm Gordeladze
\\n\\nChapter 6 Vitamin K2 and Bone Health by Niels Erik Frandsen and Jan Oxholm Gordeladze
\\n\\nChapter 7 Vitamin K2 and its Impact on Tooth Epigenetics by Jan Oxholm Gordeladze, Maria A. Landin, Gaute Floer Johnsen, Håvard Jostein Haugen and Harald Osmundsen
\\n\\nChapter 8 Anti-Inflammatory Actions of Vitamin K by Stephen J. Hodges, Andrew A. Pitsillides, Lars M. Ytrebø and Robin Soper
\\n\\nChapter 9 Vitamin K2: Implications for Cardiovascular Health in the Context of Plant-Based Diets, with Applications for Prostate Health by Michael S. Donaldson
\\n\\nChapter 11 Vitamin K2 Facilitating Inter-Organ Cross-Talk by Jan O. Gordeladze, Håvard J. Haugen, Gaute Floer Johnsen and Mona Møller
\\n\\nChapter 13 Medicinal Chemistry of Vitamin K Derivatives and Metabolites by Shinya Fujii and Hiroyuki Kagechika
\\n"}]'},components:[{type:"htmlEditorComponent",content:'This book serves as a comprehensive survey of the impact of vitamin K2 on cellular functions and organ systems, indicating that vitamin K2 plays an important role in the differentiation/preservation of various cell phenotypes and as a stimulator and/or mediator of interorgan cross talk. Vitamin K2 binds to the transcription factor SXR/PXR, thus acting like a hormone (very much in the same manner as vitamin A and vitamin D). Therefore, vitamin K2 affects a multitude of organ systems, and it is reckoned to be one positive factor in bringing about "longevity" to the human body, e.g., supporting the functions/health of different organ systems, as well as correcting the functioning or even "curing" ailments striking several organs in our body.
\n\nChapter 1 Introductory Chapter: Vitamin K2 by Jan Oxholm Gordeladze
\n\nChapter 2 Vitamin K, SXR, and GGCX by Kotaro Azuma and Satoshi Inoue
\n\nChapter 3 Vitamin K2 Rich Food Products by Muhammad Yasin, Masood Sadiq Butt and Aurang Zeb
\n\nChapter 4 Menaquinones, Bacteria, and Foods: Vitamin K2 in the Diet by Barbara Walther and Magali Chollet
\n\nChapter 5 The Impact of Vitamin K2 on Energy Metabolism by Mona Møller, Serena Tonstad, Tone Bathen and Jan Oxholm Gordeladze
\n\nChapter 6 Vitamin K2 and Bone Health by Niels Erik Frandsen and Jan Oxholm Gordeladze
\n\nChapter 7 Vitamin K2 and its Impact on Tooth Epigenetics by Jan Oxholm Gordeladze, Maria A. Landin, Gaute Floer Johnsen, Håvard Jostein Haugen and Harald Osmundsen
\n\nChapter 8 Anti-Inflammatory Actions of Vitamin K by Stephen J. Hodges, Andrew A. Pitsillides, Lars M. Ytrebø and Robin Soper
\n\nChapter 9 Vitamin K2: Implications for Cardiovascular Health in the Context of Plant-Based Diets, with Applications for Prostate Health by Michael S. Donaldson
\n\nChapter 11 Vitamin K2 Facilitating Inter-Organ Cross-Talk by Jan O. Gordeladze, Håvard J. Haugen, Gaute Floer Johnsen and Mona Møller
\n\nChapter 13 Medicinal Chemistry of Vitamin K Derivatives and Metabolites by Shinya Fujii and Hiroyuki Kagechika
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I am also a member of the team in charge for the supervision of Ph.D. students in the fields of development of silicon based planar waveguide sensor devices, study of inelastic electron tunnelling in planar tunnelling nanostructures for sensing applications and development of organotellurium(IV) compounds for semiconductor applications. I am a specialist in data analysis techniques and nanosurface structure. 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After obtaining a Master's degree in Mechanical Engineering, he continued his PhD studies in Robotics at the Vienna University of Technology. Here he worked as a robotic researcher with the university's Intelligent Manufacturing Systems Group as well as a guest researcher at various European universities, including the Swiss Federal Institute of Technology Lausanne (EPFL). During this time he published more than 20 scientific papers, gave presentations, served as a reviewer for major robotic journals and conferences and most importantly he co-founded and built the International Journal of Advanced Robotic Systems- world's first Open Access journal in the field of robotics. Starting this journal was a pivotal point in his career, since it was a pathway to founding IntechOpen - Open Access publisher focused on addressing academic researchers needs. Alex is a personification of IntechOpen key values being trusted, open and entrepreneurial. 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