Showing world production of phosphate in 2015 and 2016.
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Released this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\\n\\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
\\n"}]',published:!0,mainMedia:null},components:[{type:"htmlEditorComponent",content:'IntechOpen is proud to announce that 179 of our authors have made the Clarivate™ Highly Cited Researchers List for 2020, ranking them among the top 1% most-cited.
\n\nThroughout the years, the list has named a total of 252 IntechOpen authors as Highly Cited. Of those researchers, 69 have been featured on the list multiple times.
\n\n\n\nReleased this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\n\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
\n'}],latestNews:[{slug:"stanford-university-identifies-top-2-scientists-over-1-000-are-intechopen-authors-and-editors-20210122",title:"Stanford University Identifies Top 2% Scientists, Over 1,000 are IntechOpen Authors and Editors"},{slug:"intechopen-authors-included-in-the-highly-cited-researchers-list-for-2020-20210121",title:"IntechOpen Authors Included in the Highly Cited Researchers List for 2020"},{slug:"intechopen-maintains-position-as-the-world-s-largest-oa-book-publisher-20201218",title:"IntechOpen Maintains Position as the World’s Largest OA Book Publisher"},{slug:"all-intechopen-books-available-on-perlego-20201215",title:"All IntechOpen Books Available on Perlego"},{slug:"oiv-awards-recognizes-intechopen-s-editors-20201127",title:"OIV Awards Recognizes IntechOpen's Editors"},{slug:"intechopen-joins-crossref-s-initiative-for-open-abstracts-i4oa-to-boost-the-discovery-of-research-20201005",title:"IntechOpen joins Crossref's Initiative for Open Abstracts (I4OA) to Boost the Discovery of Research"},{slug:"intechopen-hits-milestone-5-000-open-access-books-published-20200908",title:"IntechOpen hits milestone: 5,000 Open Access books published!"},{slug:"intechopen-books-hosted-on-the-mathworks-book-program-20200819",title:"IntechOpen Books Hosted on the MathWorks Book Program"}]},book:{item:{type:"book",id:"9236",leadTitle:null,fullTitle:"Cheminformatics and its Applications",title:"Cheminformatics and its Applications",subtitle:null,reviewType:"peer-reviewed",abstract:"Cheminformatics has emerged as an applied branch of Chemistry that involves multidisciplinary knowledge, connecting related fields such as chemistry, computer science, biology, pharmacology, physics, and mathematical statistics.The book is organized in two sections, including multiple aspects related to advances in the development of informatic tools and their specific use in compound structure databases with various applications in life sciences, mainly in medicinal chemistry, for identification and development of new therapeutically active molecules. 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Upon mild to moderate hepatocellular injury or depletion, hepatocytes self-duplicate to restore the liver mass. However, when there is a massive cell loss or a continuous damage to mature hepatic cells, overwhelming the replicative capacity of the remaining hepatocytes, expansion of immature-like cells is observed at the interface between the portal area and the parenchyma in a process called ductular reaction (DR). Expression of biliary markers is a hallmark of DR cells, but nevertheless, DR constitutes a heterogeneous population of proliferating cells ranging from immature stem-like cells to more committed cells with an intermediate hepatobiliary phenotype [1, 2, 3, 4]. Cells of the DR are also called liver progenitor cells (LPC) as they have been shown to differentiate into both hepatocytes and cholangiocytes lineages in culture (reviewed in Ref. [5]). In normal livers, no DR are usually observed and LPC are seen, in two-dimensional tissue sections, as single cells located mainly in the canal of Hering, which represents the connection between the smallest ramifications of the biliary tree and the hepatocyte canalicular system [6, 7]. DR/LPC and biliary cells cannot strictly be distinguished at the histological level but based on their location and morphological differences [8]. In a three-dimensional viewpoint, DR and the biliary tree constitute together a contiguous heterogeneous epithelial structure [9]. In humans as in rodents, the histological and morphological patterns of DR vary according to injurious settings and their lineage commitment toward hepatocytes or cholangiocytes has been related to the primary site of cell loss or dysfunction [10].
Over the past decade, there has been a considerable interest in understanding DR/LPC biology. LPC are indeed seen as a potential reservoir for mature hepatocytes. Understanding the nature and differentiation process of LPC may generate cells for liver-cell therapy, which is increasingly under demand due to organ shortage for liver transplantation. Moreover, DR has also been postulated to trigger portal fibrosis [11]. Unraveling the potential mediators of DR could therefore be of great interest to modulate progression of profibrogenic reaction observed in many chronic liver diseases.
Several rodent models of liver injury associate with a DR and are instrumental to study the LPC response and its implication in liver regeneration and wound healing. These models, as in human liver diseases, exhibit a large variety of DR/LPC patterns with different morphological features, kinetics of response, and differentiation potential. The models of liver injury with DR generally combine the damage and loss of epithelial cells (hepatocytes and/or cholangiocytes) with the inhibition of the proliferative capacity (replicative senescence) of mature epithelial cells. Toxins [12, 13], carcinogens [14, 15], or modified diets [16, 17] have been used to induce cell injury, either alone or associated with surgical removal of part of the liver to amplify liver cell depletion. Ethionine, 2-acetylaminofluorene (AAF), and retrorsine are used to block the ability of mature epithelial cells to divide and prevent them from contributing to the liver regeneration process. In mice, dietary manipulations are regarded as convenient, efficient, and reproducible models to induce a robust DR, without need for animal handling, repeated injections, or surgical manipulation. The two most popular dietary DR models are a choline-deficient diet supplemented with ethionine in the drinking water (CDE) or a diet enriched in 3,5-diethoxycarbonyl-1,4-dihydrocollidine (DDC).
In the literature, the DDC and CDE models are often used equivalently to study the LPC response and their role in tissue repair. However, DR in those two models exhibits major etiological and phenotypical differences. Such differences recapitulate the specificity of the pathophysiological responses to distinct injurious processes. DR activation, expansion, and capacity for differentiation are dictated by the nature of the cellular injury and by the differential microenvironment changes.
In this chapter, we will first describe the pathophysiological mechanisms at play in each model and the experimental procedures to induce DR with the CDE or DDC diet. A description of the hepatic lesions in terms of the cellular compartment injured after CDE and DDC feeding and highlight of the unique character of each model with regard to the DR phenotype, proliferation, lineage commitment, and microenvironment will be explored. Finally, the relevance of these models to study and understand the diversity of DR seen in human chronic liver diseases will be addressed.
The CDE model consists of ad libitum administration of a choline-deficient diet together with procurement of ethionine in the drinking water. Choline is provided by food intake and contributes to the structural integrity and signaling function of cell membranes. A choline withdrawal leads to a decreased synthesis of phosphatidylcholine, a phospholipid crucial for cell membrane and a major building stone of the very low-density lipoprotein particles produced by hepatocytes to export triglycerides. Choline deficiency causes intracytoplasmic fat accumulation, hepatocyte dysfunction, and cell damage [18, 19]. Such (extensive) hepatocellular damage results in high hepatocyte replication ratio, causing their exhaustion and restraining the production of hepatic drug metabolism-related enzymes [20]. Ethionine, a synthetic amino acid, specifically targets the hepatocytes in which, when provided in large excess, it competes with its naturally occurring analog methionine. Competition of ethionine with methionine favors the synthesis of S-adenosyl ethionine (SAE) instead of S-adenosyl methionine (SAM). Consequently, an ethyl group is transferred instead of a methyl group in methylation reactions hereby generating abnormal proteins, lipids, RNA, and DNA molecules, which results in hepatocytic cell damage [21]. Prolonged feeding with ethionine produces liver tumors with extensive LPC proliferation [22]. However, administration of ethionine in supplement to a choline-deficient diet greatly shortens the time required for LPC proliferation [22]. Although, the exact mechanism of action of CDE-induced injury is not well known, it appears that the combined administration of ethionine with choline-deficient chow induces a liver injury in which the hepatocytes are specifically targeted and the replication of the surviving hepatocytes is inhibited [23]. Hepatocyte proliferation to replace damaged liver cells is prevented and activation of the LPC compartment ensues. Several publications characterized the kinetics of the LPC response and liver damage to CDE [24, 25, 26, 27]. Briefly, short-term CDE feeding results in steatosis, inflammation, LPC expansion (DR), and fibrosis that progress in parallel. Cirrhosis and hepatocellular carcinoma may be observed in long-term studies. We intend to describe and analyze in depth the morphology and differentiation capacity of DR after 3 weeks of CDE (except when specified otherwise), at a time when pathological damages are installed and DR robustly established.
After 3 weeks, CDE livers are pale with signs of steatosis throughout the parenchyma. Liver weight is comparable or slightly lower than the deep-red control livers (Figure 1A and B). Signs of hepatocellular injury are observed with necrotic and apoptotic hepatocytes while bile ducts appear normal within the portal triad (Figure 1G) [23, 28]. Also, serum alanine amino-transaminases are increased while bilirubin levels are in the near normal range, indicative of hepatocytic damage (Figure 1D and E).
Pathophysiological mechanisms and DR phenotype in the CDE and DDC models. Livers retrieved from control mice (A), mice receiving the CDE (B), or DDC diet (C) for 3 weeks. Serum biochemical measurements for total aminotransferase (ALT) (D) and bilirubin (E) showed increased ALT and near normal bilirubin in the CDE model and increased serum bilirubin with slightly elevated ALT levels in DDC, indicative of hepatocellular damage upon CDE treatment and of primarily biliary injury after DDC diet. Liver sections stained with anticytokeratin 19 (CK19) in control (F), CDE (G), and DDC (H) livers after 3 weeks of diet. In control, CK19 staining reveals bile ducts and LPC as isolated cells close to the periportal tract. In CDE livers, besides bile ducts, DR CK19+ cells are strongly increased in number, forming cells organized in filaments expanding inside the lobule. After DDC feeding, in addition to the larger preexisting bile ducts, CK19+ newly formed DR structures are composed of small cuboidal cells, irregular in size and shape accumulated around the portal area. Plastination of the bile duct system reveals delicately structured biliary tree in control mice (I), a denser biliary network after CDE feeding (J), and dilatation of intrahepatic bile ducts in DDC-fed mice (K).
In the CDE model, DR expansion, seen on two-dimensional (2D) sections by staining with a biliary marker such as cytokeratin (CK) 19, is observed arising from the portal area and invading progressively the parenchyma (Figure 1G). First observable after approximately 1 week of CDE feeding, the DR progressively amplifies to a maximum around 3–4 weeks [23, 24, 29]. DR cells are small cells with a high nuclear-to-cytoplasm ratio, usually uniform in size with a fusiform shape and oval nuclei. On 2D liver sections, they are found as individual cells, grouped in multifocal clusters or organized in a single or double row of cells forming arborizing structures (Figure 1) [24]. Architectural three-dimensional (3D) analysis of the biliary tree remodeling in response to CDE reveal that DR are connected to the preexisting bile ducts and that biliary branches intricately split around the portal vein with a random directionality [9]. Moreover, plastination of the bile duct system reveal a denser biliary network after CDE feeding (Figure 1J).
Finally, with regard to LPC capacity of differentiation in vivo, DR cell-tracking experiments using different transgenic mouse models [23, 28, 30] indicate that, upon CDE diet, a small number of DR cells do differentiate into hepatocytes: in the process LPC lose biliary markers, grow in size, and acquire mature hepatocyte morphological features and functional proteins. Although differentiation is consistently reported, only few DR-derived hepatocytes are reported in this model (<2.5% of hepatocytes).
Although being widely used, the CDE model is difficult to handle and researchers are confronted with difficulties and ethical issues due to variability in the LPC response, well-being of the animals, morbidity, and mortality. Here, we will review several factors influencing the LPC response to the model. These parameters must be taken into account and controlled to strengthen the model and provide reproducibility.
Rodent food manufacturers can easily provide food in which the choline content is strictly controlled. Although low choline dietary content could be used [31], we will describe here a model using dietary choline deficiency. The second parameter to adjust for is ethionine supplementation. In the literature, the amount of ethionine in the water varies from 0.05 to 0.165% (wt/vol). Mice are not fond of ethionine (due to bad smell), and usually decrease water consumption. This makes it difficult to control effective ethionine intake. Addition of 5% sucrose, choline-free orange juice, or fruit syrup is sometimes used to increase the attractiveness of the drink and this is most of the time not reported in the experimental protocol [9, 23, 28, 29, 32, 33, 34]. Ethionine smell increases with exposure to the ambient air and we found that we could maintain stable water intake by replacing ethionine-containing water by a fresh solution every day. Although this sounds trivial, control over ethionine solution consumption is crucial as both variation in ethionine intake and (severe) dehydration may influence LPC response and induce large interindividual variation in the model. To circumvent this, Passman et al. also propose to include ethionine in the chow [29].
LPC response and morbidity vary according to weight and age of the mice at the time of introduction of the CDE diet. Mice above 25–30 g will be quite resistant to the diet and if they are too old, perhaps because of loss of cell plasticity, LPC response will be discrete. In parallel, if they are too young or too little (<15 g) at the time of dietary exposure, toxicity and ensuing mortality might be excessively high. With the administration of the CDE diet to mice of 6 weeks of age and with a body weight between 18 and 20 g, we and others show a substantial and reasonably reproducible LPC response while maintaining the well-being of the animals [24, 28, 29, 35]. Mice are experiencing the most severe effects of the diet during the first week of administration. Following the first few days, significant weight loss is observed, often associated with mortality [29, 36]. Approximately 1 week after the onset of the CDE treatment, the mice adapt, regain weight, and show (normal) physical activity and behavior. Thus, by respecting the simple rules proposed above, body weight loss may be limited to 10% of the starting body weigh during the first week with weight stabilization thereafter. Importantly, sensitivity to the dietary regimen and magnitude of liver damage and LPC reaction largely vary according to the genetic background of the mice [17]. This imposes the use of an appropriate control group (best being littermates) when comparing the effect of gene deletion or addition in genetically modified animals.
The DDC model consists of ad libitum administration of a diet enriched with the porphyrinogenic agent 3,5-diethoxycarbonyl-1,4-dihydrocollidine (DDC) with normal water. Exposure to a DDC diet provokes the inhibition of the mitochondrial enzyme ferrochelatase, catalyst of the insertion of ferrous iron into protoporphyrin IX to form heme, leading to progressive accumulation of protoporphyrin. This brown pigment first accumulates in the cytoplasm of parenchymal cells and in Kupffer cells. Because of its hydrophobic nature, the excess of protoporphyrin can only exit the liver through biliary excretion, leading to precipitation of this poorly soluble molecule in bile canaliculi and bile ducts, forming crystals increasing in size and number [37]. After 3 weeks of DDC diet, accumulated pigments plug and obstruct the lumen of the smaller branches of the biliary tree and confer a dark coloration to the liver (Figure 1C). Bile ducts, usually recognizable as monolayer rings of small cuboidal cholangiocytes delineating a central lumen, show profound morphological alterations while hepatocytes have a normal appearance except for pigment coloration (Figure 1H). This indicates that the DDC dietary regimen mostly damage the biliary system, which is additionally supported by increased serum bilirubin (Figure 1E). We observed moderately elevated transaminases levels (two to threefold time, Figure 1D) although another group reports higher transaminase levels after DDC feeding [16]. In later stages, livers in DDC-fed mice develop pericholangitis and periductal onion skin-like fibrosis. Our discussion here analyzes DR morphology and microenvironment after 3 weeks of DDC feeding.
In the DDC livers, bile duct damage is associated with a biliary response in which dysmorphic cholangiocytes proliferate in the portal area. In all portal tracts, DR expands as multiple small pseudo-ducts arising next to the larger preexisting bile ducts (Figure 1H). These newly formed ductular structures are composed of small cuboidal or more cylindrical cells, irregular in size and shape, assembled in tube-like structures outlining a lumen in most cases, sometime plugged by porphyrin crystals. In contrast to infiltrating DR in CDE livers, DR expansion observed in DDC livers remains enclosed within the portal mesenchyme. No parenchymal invasion crossing the boundaries of portal mesenchyme was observed nor did those reactive cells, always observed as a cluster, adopt a phenotype supporting migration. However, the portal mesenchyme extends and may bridge distant portal spaces (Figure 1H). 3D biliary analysis of DDC livers identifies branches randomly directed around the portal vein, connected to the biliary tree but forming apparent distinct structures from the large-diameter bile ducts [9]. Moreover, 3D plastination of the DDC-fed mouse confirms slight focal dilatation of intrahepatic bile ducts and porphyrin plugs while biliary network seems to be less dense [16] (Figure 1K).
Finally, concerning LPC capacity of differentiation, upon DDC-induced injury, there is no evidence that cells of the neo ducts undergo hepatocytic cell differentiation [23, 28, 30]. When animals are reversed to a standard chow after DDC diet, the degree of DR expansion decreases, but still with no evidence that DR cells differentiate into hepatocytes. Because a specific LPC marker, that is, exclusively expressed in LPC and not in cholangiocytes, is lacking, we are currently unable to experimentally address the contribution of LPC to biliary regeneration in vivo during disease evolution. We can, however, hypothesize that, if not entirely supported by proliferation of mature cholangiocytes, LPC located at the most proximal part of the biliary tree contribute to neo duct formation in the DDC model [9].
Contrasting with the CDE model, the DDC model is robust and reproducible and has little impact on animal welfare. In all studies, diet (standard rodent chow) is supplemented with 0.1% (wt/wt) of DDC. Similarly, the different mouse strains tested so far develop comparable hepatic phenotype to DDC feeding [16] although differences in susceptibility and kinetics of the response might be expected according to strains. Of note, DDC diet applied to rats does not induce any LPC response [38].
The literature brings every day new evidence that the orchestrated interplay between proliferating hepatocytes or cholangiocytes, extracellular matrix-producing myofibroblasts, inflammatory cells (such as macrophages, neutrophils, or lymphocytes), and endothelial cells is pivotal in the regulation of DR expansion and differentiation. We will thus compare the microenvironment accompanying DR in the CDE versus DDC model.
Extracellular matrix and collagen deposition associates with DR. In the CDE model, a thin and loose web of collagen fibers is associated with invading DR cells, while collagens in DDC livers thicken the portal mesenchyme and abundant extracellular matrix accumulates in clots or thick concentric layers around the neo-formed pseudo-ductular structures (Figure 2A–D).
Comparison of the extracellular matrix deposition and the myofibroblast expansion between the CDE and DDC models. Liver sections obtained from mice receiving the CDE (A, C, E) or DDC diet (B, D, F) were stained with Sirius red to highlight fibrillary collagen (A-D), or αSMA and CK19 expression (E and F). In CDE livers, a collagen meshwork covers the whole parenchyma (A) with fibers elongating from the portal area into the lobule (C). In DDC livers, collagen fibers accumulate around the portal area to shape the portal mesenchyma (D), delimiting the boundaries of DR (F). At lower magnification, portal-portal bridging is observed (B). αSMA+ myofibroblasts show a distribution pattern similar as the collagen deposition. αSMA+ myofibroblasts infiltrate the lobule, chaperoning CK19+ DR cells in the CDE model (E), while in the DDC model, αSMA+ myofibroblasts rather accumulate concentrically around the DR structures (F).
The localization of myofibroblasts, the cells chiefly involved in matrix synthesis and remodeling, adopted a pattern similar to that of the collagen deposition in both models, meaning that DR is at all times associated with myofibroblasts. In CDE livers, myofibroblasts chaperone the DR cells while they penetrate deep into the liver lobule (Figure 2E). Conversely, in the DDC model, myofibroblasts densely populate the portal mesenchyme and accumulate rather concentrically around DR (Figure 2F).
Laminin is a component of the basal membrane delineating the basal pole of cholangiocytes. Basement membrane is essential to establish the cholangiocytes polarity and to support a tubular structure with a lumen [39]. By contrast, hepatocytes do not lie on a basement membrane. In the CDE model, DR is anchored onto a laminin-rich basal membrane intermingled with collagen. This layer of laminin has been proposed to maintain the immature/biliary phenotype of DR cells and to provide a holding structure facilitating migration of DR into the lobular parenchyma in the CDE model [28, 40]. Moreover, decreased density of laminin and extracellular matrix in CDE livers is associated with enhanced hepatocytic differentiation of DR cells [28]. Indeed, during DR differentiation process, DR cells progressively lose contact with the laminin-rich basement. And when animals are reversed to a standard chow (supply of choline and cessation of ethionine administration) after CDE exposure, the injury reverses, DR, extracellular matrix, and laminin deposition progressively lessen, and concomitantly the number of DR-derived hepatocytes increases.
In DDC livers, laminin deposits as thin basal membrane outlining the DR in a pattern similar to that seen around normal bile ducts, with collagen stacked as separate sheets encircling newly formed DR.
In response to liver injury, Kupffer cells, the hepatic macrophages, activate and participate to the recruitment of the inflammatory reaction. In CDE livers, enlarged and proliferative Kupffer cells are strongly associated with invading DR within the parenchyma while no portal inflammation is observed [41]. DDC-induced proliferation of the ducts is accompanied by a dense macrophage and neutrophil granulocytic infiltrate around small and larger bile ducts, further supporting that biliary structures are first concerned by the injurious and healing responses in this model [16].
Kupffer cells do not influence DR expansion but modulate its invasive behavior and its specification, through modulation of the density of extracellular matrix as well as via Notch and Wnt signaling pathways [41, 42]. Numb, a direct transcriptional target of Wnt and a negative regulator of Notch, is downregulated in LPC during biliary regeneration, promoting biliary specification via the Notch pathway. While during hepatocyte regeneration, macrophage-derived canonical Wnt signaling maintains Numb within LPC and Notch signaling is reduced, promoting hepatocyte specification [42].
Experiments performed in lymphocyte-deficient mice fed on CDE suggest that natural killer cells and T-cells participate also to LPC expansion, presumably through their proinflammatory cytokine production [43]. Moreover, TNF-like weak inducer of apoptosis (TWEAK), produced by T-cells and activating its receptor fibroblast growth factor-inducible 14 (Fn14), is suggested to be an exclusive LPC mitogen. After both CDE and DDC treatment on Fn14 knockout mice, a significant reduction of the LPC response is observed [31, 44].
As described above, DR requires a typical niche provided by extracellular matrix-producing and inflammatory cells, which are located in the sinusoids closely adjacent to DR. Additionally, sinusoidal endothelial cells themselves could also have an important role in regulating DR. Signaling molecules specifically expressed within the endothelial compartment of the central vein have been shown to have a crucial role in liver zonation [45]. Moreover, in another model of liver injury, hepatocytes divide along the closest microvessel as order principle to restore liver architecture [46]. Either a signaling or a guiding role of endothelial cells on LPC response could be envisaged. However, so far, no experiments have been done to study endothelial regulation of DR in the CDE or DDC model.
In humans, DR is seen in most chronic liver injury, irrespective of the etiology. Historically, DR has been categorized on morphology into “typical” and “atypical” DR, based on rodent studies [47]. Typical DR have a lumen lined by cuboidal cells and are the result of proliferation of preexisting ductules, in analogy with the DR seen after biliary obstruction, while atypical describes thin, elongated structures that extend into the lobules and lack discernible lumen as preferentially seen after hepatocytic damage. Therefore, DDC diet best models typical DR, while CDE diet replicates pathological pattern of atypical DR. However, this dichotomic classification was discouraged some years ago because it could not readily accommodate the range of patterns seen clinically [10]. Another classification schemes attempted to integrate the histologic features, inciting disease and immunophenotyping of DR [48, 49]. However, not all DR fit in this classification, and especially not when biliary obstruction becomes chronic (as in the DDC livers). So far, there is a lack of consensus regarding DR classification in humans, as DRs are diverse, covering a spectrum of features rather than clear subphenotypes [10].
However, based on histological analysis, the DR phenotype in the CDE model resembles the one observed in human chronic Hepatitis C virus (HCV) infection depicting portal fibrosis and in a series of autoimmune hepatitis (AIH) (Figure 3A–D). HCV- and AIH-associated DRs have only a vague or no lumen, and comprise small elongated cells with little cytoplasm extending in the periportal parenchyma and associated with dense collagen fibers [10, 11, 50]. In HCV, the extension of DR into the parenchyma and DR severity correlate with the severity of fibrosis and the inflammatory activity, supporting that extracellular matrix and inflammatory signals influence DR [50]. DR in AIH has been proposed to represent a regenerative response as DR persists after the inflammatory activity subsided following immunosuppressive treatment [51]. At the early stage of fibrosing, cholestatic variants of coinfection with hepatitis B and C, expanded DR into the hepatic parenchyma also resemble the DR phenotype seen in CDE livers [10, 52]. With regard to LPC fate, the observation of a phenotypic continuum between DR cells and hepatocytes in the livers of patients suffering from HCV supports differentiation of LPC toward hepatocytes [4, 50]. Besides hepatitis, the CDE diet also recapitulates features of the DR associated with lipid accumulation (steatosis) as in chronic alcoholic and nonalcoholic fatty liver diseases [11, 29, 42, 53].
DR observed in human chronic hepatitis C infection and in primary sclerosing cholangitis. Liver stained with anticytokeratin 7 (CK7) and Masson-trichrome of an HCV case with mild inflammation (A and B), AIH with moderate inflammation (C and D), and PSC with cholangitis, edema, and portal fibrosis (E and F). HCV- and AIH-associated CK7+ DRs (A and C) have only a vague or no lumen and comprise and elongated cells with little cytoplasm extending in the periportal parenchyma and associated with dense collagen fibbers (B and D). While the CK7+ ductular proliferation seen in PSC (E) is enclosed in portal mesenchyma and concentric periductular fibrosis occurs (F).
The DR pattern seen in DDC livers is more comparable to that of chronic fibrosing cholangitis such as primary sclerosing cholangitis (PSC) and primary biliary cholangitis (PBC) with DR proliferation restricted within the portal area and accompanied by concentric periportal fibrosis (Figure 3E and F). In these diseases as in DDC, the primary damage is directed toward cholangiocytes. Intrahepatic bile duct destruction and ductopenia seen in advanced PBC and the fibrous obliterative lesions of PSC do not occur in the DDC model, a phenomenon most likely related to the specific immune component of PBC and PSC which is lacking in the DDC model.
As mentioned above, the Notch and Wnt signaling pathways are involved in the divergence of DR cells fate toward hepatocyte or biliary cells observed in response to CDE versus DDC. Similarly, in human diseases, prevalence of Notch signaling, driving biliary phenotype, is strong in PSC while the expression of Numb, a negative regulator of Notch, is more elevated in HCV samples compared to PSC [54]. Moreover, β-catenin, a component of the Wnt pathway, is found within the cytoplasm and nucleus of human DR cells of HCV-infected livers, signing enhanced Wnt signaling and promoting hepatocyte regeneration, whereas in PSC, β-catenin is predominantly localized to the cell surface, suggesting low activation of the canonical Wnt signaling pathway promoting biliary regeneration [42].
In summary, the CDE diet targets specifically hepatocytes and induce DR-containing elongated cells of an undifferentiated and migration-supporting phenotype expanding from portal tracts into the parenchyma. Myofibroblast activation and extracellular matrix deposition precedes this cell expansion, and a laminin-rich sheet sustains those DR while macrophages associate with invading DR. In the DDC model, accumulating protoporphyrin obstructs the hepatobiliary system leading to biliary damage and resulting in highly proliferative cells forming bile duct-like structures remaining restricted to portal mesenchyme, delineated by a thin layer of laminin and accompanied by dense portal inflammation. Moreover, cell-tracking experiments revealed that DR cells are able to generate a small number of functional hepatocytes after CDE but not after DDC exposure. Finally, the DR phenotype and signaling pathways involved in LPC differentiation in the CDE model mirrors the one observed in chronic HCV infection presenting signs of fibrosis and autoimmune hepatitis, while the DDC model could be used to study biliary injury such as PSC or PBC in humans. We believe that characterization of the most widely used dietary DR mouse models will help our understanding of the diversity of DR patterns observed in humans and will help the researchers to select the appropriate model in relation to the specific question addressed.
0.12% of earth crust is made up of phosphorus mineral. P is a nonrenewable natural resource present in all types of rock and soils, in all living cells, and however can form complex compounds. Mineral deposits are the major supply of phosphorus. All phosphate mineral was derived from apatite by weathering. Mostly phosphate is found in different forms like quartz, calcite, dolomite, apatite, Fe-oxide minerals and clay minerals. Apatite mineral is used for manufacturing fertilizers. Extraction of phosphorus depends on the physical properties of the rocks and its geological setting.
Since ancient times man used natural resources such as manures, vegetables material, and bones as fertilizers. In 1840 Liebig, the German chemist, suggested the formation of superphosphate by dissolving bones in sulfuric acid that made the P more available to plants. This practice becomes so popular that bone supply is restricted in a very short time. To overcome this problem, some workers started extraction of phosphorus from rocks; in 1847 the first commercial production of P rocks from the mining of coprolites began in Suffolk in Great Britain and peaked in 1876 when about 25,000 metric tons were mined.
Presently all the Phosphate reserves that are found all over the world are not “mineable” deposits, as mining of them are not economically feasible. The United States is the highest phosphate-producing country in the world, while Morocco and China are the second and third countries with respect to phosphate production. Australia and Canada are recently known sites of phosphorus mining. There are rich deposits of phosphate found in Mongolia and Peru that will fulfill the need in the future. Florida phosphate industry becomes one of the major producer and exporter of phosphate fertilizer due to good transportation and industrial infrastructure facility in America and also because a substantial layer of phosphate is only 15 to 50 feet below a soft overburden. The phosphate mining in Central Florida overshadowed other sources because of low cost of mining, large deposits and the good quality of phosphate content of Florida rock. Florida is presently providing approximately 75% of the nation’s supply of phosphate fertilizer and about 25% of the world supply. In 2000, mining operations began in Ontario, Canada, of North America. Florida’s phosphate is part of a deposit that stretches across the state and up the coast to the Chesapeake Bay. The phosphate mining is expanded from Central Florida to Polk and Hillsborough counties, south, to Hardee, DeSoto, Manatee and Sarasota counties. In Northern Florida phosphate deposits are present in Hamilton, Columbia and Suwannee counties.
Some three decades ago in 1880, Dr. C. A. Simmonsin of England, who owned a rock quarry for building stone in Hawthorne, sends some of his rocks to Washington, DC, for analysis. The analysis determined the presence of phosphate in the rock samples, and in 1883 he made the first attempt for mining phosphate in Florida. But it was in 1889 by Albertus Vogt and others in Marion County who began the production of the first hard rock by the Marion Phosphate Company. This was later in 1890 followed by the Dunnellon Phosphate Company, in which Vogt had ownership interest, and in this way the area was flooded by thousands of prospectors, and the great Florida phosphate boom had begun. By 1894 more than 215 phosphate mining companies were operating statewide. The boom brought wealth. But in 1900 due to consolidation and capitalization, this number had dwindled to about 50. In 1881, Captain J. Francis LeBaron, chief engineer of the US Army Corps, during his survey of Peace River of Polk County, analyzed river pebbles and confirmed the presence of phosphate, but at that time this discovery did not catch much attention. In 1886 John C. Jones and Captain W. R. McKee, of Orlando, discovered high grade phosphate along the Peace River which led to the formation of an association known as the Peace River Phosphate. Mining activity along the Peace River proceeded both in the river itself and on the adjacent land. The so-called river pebble mining was the first to be exploited. In 1888, the first shipment of Peace River phosphate pebble was launched by Arcadia Phosphate Company about a year ahead of the Peace River Phosphate Company. This phosphate discovery was kept relatively quiet. Rumors of no phosphate in Central Florida spread as a result; Polk County’s phosphate deposit took a back seat the first 15 years to the hard rock region to the north. The Florida Phosphate Company and the Pharr Phosphate Company were the two phosphate mining plants found in pebble district till 1890. In 1891 Pharr started shipment of land pebble for the first time; due to this there occurred a boom in the rate of river pebble production in 1893. Phosphate mining came to North Florida in the 1960s when Occidental Petroleum Company was looking for a way to get into the fertilizer business to get profit. Occidental went north and opened a mine in White Springs where it mined phosphate until 1995, when the Potash Corporation of Saskatchewan (PCS) purchased the operation. Nowadays Mosaic and PCS Phosphate, White Springs, are the two phosphate mining companies in Florida, and the third one are US Agri-Chemicals which produce phosphate fertilizers in Central Florida.
A blanket of phosphate deposits covers much of the Peninsular Florida has a large phosphate deposits which consists of approximately equal parts phosphate rock, clay and sand, averages 12 to 15 feet in thickness. The matrix is buried beneath a soil that is 15–30 feet deep. By the end of 1999, approximately 300,000 acres of land, or more than 460 square miles, had been mined in Florida. Polk County is the heart of the Bone Valley mining region, and the mineable deposit in this area stretches to Hillsborough, Hardee, Manatee, and DeSoto counties. The large depositions were also found in mining in North Florida’s Hamilton County from a mineable area that extends into Columbia and Suwanee counties. Similar deposition is found in both the areas. Mining in Central Florida has been moving south. As sites mine out, the draglines move to where the contiguous deposit of phosphate pebble is found. Toward the south the quality of rock decreases which brings technological challenges for the mining industries. During the past years, mining is slowed down in Polk County’s southern fringe. In 2000 closing of IMC Clear Springs and Noralyn mines conveyed a close to active. Currently phosphate mining companies has opened new mining sites in Manatee, DeSoto and Hardee counties.
The fertilizer that quickly became the item of commerce as most widely used by the growers today, and it had the highest concentration of phosphate and nitrogen at 18 N–46P2O5–0K2O.
This fertilizer is essentially the same as DAP, but it has a lower concentration of nitrogen at 11 N–52P2O5–0K2O. It is completely water soluble and has granular material; it mixes well and frequently serves as an ingredient in bulk-blended fertilizers.
It is very similar to the superphosphate fertilizer that provides 46% P2O5, some calcium and sulfur to plants. GTSP is formed by reaction of phosphate rock with phosphoric acid.
It’s an acid used to make a concentrated or fluid fertilizer. PCS is the acid produced only by Florida Company in North Florida.
Phosphoric acid is used in granulation plants where ammonia is added to phosphoric acid to produce the ammoniated phosphate fertilizer. Purified food-grade phosphoric acid is used in making soft drinks.
Defluorinated phosphate rock or phosphoric acid is used to make animal feed supplements by combining phosphate rock with phosphoric acid, sodium carbonate and then calcine or react it with lime to get dicalcium phosphate.
Sulfuric acid
This acid is used to produce phosphoric acid after reacting with phosphate rocks at phosphate plants.
Sedimentary marine phosphorites are the principle resources of phosphate rock. The world’s largest sedimentary reservoirs are found in North Africa, China, the Middle East, and the United States. Valuable igneous sedimentary reservoirs are also found in Brazil, Canada, Finland, Russia, and South Africa. Substantially large phosphate deposits have been spotted near the Atlantic Ocean and the Pacific Ocean shown in Table 1 and Figure 1. World resources of phosphate rock are more than 300 billion tons. There are no imminent shortages of phosphate rock.
Phosphate rock production worldwide in 2017, by country (in 1000 metric tons).
It was observed that applications of H3PO3 and phosphite (Phi) were less effective as compared to phosphoric acid (H3PO4) and its derivatives on the first crop. With increasing rates of phosphite (Phi), phytotoxic effects were detected on the crop yield. However, nutritive role of Phi in growth response was evident when compared to the zero-P control. Whereas researchers found Phi and H3PO3 treatments beneficial to the second crop, this was due to probable conversion of Phi to phosphate in the soil. In general, better yield was obtained when Phi materials were used on soils with limestone. Further scientific studies related with the significance of H3PO3 and its salts in agriculture did not occur for nearly 30 years, but rather their performance against plant diseases was mentioned [1]. During the disease control analysis, many incidents related to the plant’s physical and chemical mobility were observed when the plants were treated with H3PO3 or its salts in the absence of plant pathogens, some of which are described below. Ouimette and Coffey [2] reported that the Phi were more readily absorbed into plant tissues than phosphates—very important in crops with leaf surfaces that resist foliar spray uptake. In a comprehensive review given by Guest and Grant [3] related with the complex action of phosphonates, several unique features of this chemical group were recounted. For example, Phi is a rapidly absorbing nutrient, which translocates from xylem to phloem according to normal source-sink relationships for nutrient element materials. Guest and Grant [3] reported that the Phi is more persistent as it metabolized slowly in plant tissue as compared with phosphate and does not participate in all the same biochemical pathways as phosphate. Adam and Conrad and Casida [4, 5] confirmed their results through experiments where bacterium Pseudomonas fluorescens 195 showed the ability to oxidize Phi and also discharge it in the growth medium as phosphate. Malacinski and Konetzka [6] repeated the same work and reported that a short adaptive period was required before oxidation of Phi by organisms, and this whole process took 14–15 weeks. Bezuidenhout et al. [7] during their study reported first time that the Phi can also be converted microbially to phosphate within plant tissues and identified three genera of bacteria (Alcaligenes, Pseudomonas and Serratia). These findings complemented the previous observations given by Rothbaum [8] that elemental P in soil was oxidized non-enzymatically under particular temperature and water. Busman et al. [9] reported that the phosphate fertilizer applied to the soil will not be utilized by the crop in the first season. Rothbaum and Baillie [8, 10] observed that Phi was less adsorbed than phosphate by the same soil. This lower ‘phosphate fixation’ improved growth of Phi-treated soil, with a period gap, as compared to phosphate-treated soil. Rhone-Poulenc Ag Company of the United States expressed concern to the US Environmental Protection Agency (EPA) about classifying a fungicide based on H3PO3 salts as a biochemical pesticide and affirmed the non-enzymatic oxidation of Phi to phosphate occur naturally over time. Lovatt [11] discovered that foliar application of K3PO3 to P-deficient citrus seedlings restored plant growth. This demonstrated that through metabolic processes, Phi was readily taken up by plant leaves and replaced phosphate as a source of Frazier and Waerstad [12] tested the composition and solubility of Phi to analyze the potential of this class of materials for increasing the plant nutrient element content of liquid fertilizers. Albrigo [13] reported the positive response of Phi on winter pre-bloom foliar of Valencia oranges which were increased flower number, fruit set and yield, plus increased total soluble solids. Additional studies by Lovatt [14] on foliar fertilization of citrus showed that application of K3PO3 in May and July to navel orange significantly increased the number of large-size fruit, total soluble solids and the ratio of soluble solids to acid, compared to control fruit. Biagro Western Sales, Inc., Visalia, CA, took the lead in commercialization of Phi-supplying fertilizer products patented by the University of California Anon and Lovatt [15, 16]. Today farmers are well educated and formed community of producers; they analyze themselves the effect of new Phi products on both soil and crop. In a practical sense, acceptance by discriminating growers is strong evidence that the benefits of H3PO3-derived fertilizers are standing up to their ultimate test—the real world of agricultural crop production.
Rock phosphate is one of the basic raw materials needed in the manufacture of phosphatic fertilizers like single superphosphate, diammonium phosphate, nitrophosphates, etc. Commercial rock phosphate occurs in nature as deposits of apatites (bearing minerals) along with other accessory minerals such as quartz, silicates, carbonates, sulfates, sesquioxides, etc. Four types of rock phosphate minerals are carbonate apatite [3Ca3(PO4)2.CaCO3], fluorapatite [3Ca3 (PO4)2.CaF2], hydroxyapatite [3Ca3(PO4)2.Ca(OH)2], and sulpho apatite [3Ca3 (PO4)2.CaSO4]. Because of their well-developed crystalline formation property, the apatites of igneous and metamorphic origin are generally regarded as less reactive. However, the apatites of sedimentary rock deposits are soft minerals possessing microcrystalline structure and are of major commercial importance for direct application in the soil [17].
The classification of reserves of indigenous rock phosphate as done by the Indian Bureau of Mines, and the purpose for which each grade can be used is given in Table 2.
Countries | Mine production | |
---|---|---|
2015 | 2016 | |
United States | 27,400 | 27,800 |
Algeria | 1400 | 1500 |
Australia | 2500 | 2500 |
Brazil | 6100 | 6500 |
China | 120,000 | 138,000 |
Egypt | 5500 | 5500 |
India | 1500 | 1500 |
Israel | 3540 | 3500 |
Jordan | 8340 | 8300 |
Kazakhstan | 1840 | 1800 |
Mexico | 1680 | 1700 |
Morocco and Western Sahara | 29,000 | 30,000 |
Peru | 3880 | 4000 |
Russia | 11,600 | 11,600 |
Saudi Arabia | 4000 | 4000 |
Senegal | 1240 | 1250 |
South Africa | 1980 | 1700 |
Syria | 750 | — |
Togo | 1100 | 900 |
Tunisia | 2800 | 3500 |
Vietnam | 2500 | 2800 |
Other countries | 2470 | 2410 |
World total (rounded) | 241,000 | 261,000 |
Showing world production of phosphate in 2015 and 2016.
Grade | P2O5 (%) | Reserve (mt) | Remarks |
---|---|---|---|
High | +30 | 15.27 | Considered for wet production of fertilizers |
Medium | 25–30 | 18.95 | Considered mainly for partially acid rock phosphate and for processed phosphates after less beneficiation |
Low | 11–25 | 55.22 | Approx. 20% P2O5 grade and relatively more reactive material may be considered for partially acidulated rock phosphate production and others for direct application |
Unclassified | 170.04 | ||
Total | 259.48 |
Classification of known reserves of indigenous rock phosphate in India.
Including all grades and types of rock phosphate, the known global resources are in the order of 163,000 million tons. Though globally adequate, rock phosphate is inequitably geographically distributed. Africa holds about 41%, the United States has 21%, former USSR 13%, the Middle East 10%, Asia 8%, and South America 3%, while Australia, New Zealand and Oceania together reported for only 2% and Europe >1%. Phosphate rock resources in India is, however, not very comfortable as it possesses a resource of only 260 million tons (0.19% of the world) of rock phosphate of all types and grades, catering the agricultural needs of 1/6 of the population of the world. Out of the total rock phosphate resource, the country has a predominance of low grade rock phosphate having only 15.27 million tons reserve of high grade rock phosphate (Table 1), and the remaining low grade rock phosphate is unacceptable to P fertilizer industry due to its very low P2O5 and high CaCO3 content [18]. The current annual domestic demand of high grade rock phosphate is of the order of 4 million tons. Out of which 95% is consumed in agriculture sector as a source of P fertilizer. The domestic production of about 1.4 million tons/year of rock phosphate could hardly meet 35% of the total demand, and the remaining (65%) demand is met through imports. P fertilizer industry largely depends on sulfur, phosphoric acid, and ammonia besides rock phosphate. India imports around 1.7 million tons of sulfur, 2–4 million tons of phosphoric acid, 1.5 million tons of NH3 and 4.9 million tons of rock phosphate for phosphate industry which constitutes a substantial part of our international trade in fertilizer raw material. Thus, the rapidly increasing price of soluble phosphatic fertilizer has raised interest in cheaper alternatives. Under such conditions, we must explore new methodologies for the utilization of indigenous low grade rock phosphate by converting it into a potential resource of P for direct application to the soil. The direct utilization of indigenous rock phosphate deposits could only alleviate the dependence of the country on foreign suppliers.
Soils has an eminent reserve of total P, but very little amount of P is actually available to the plants to support their growth to fulfill the requirement; continuous application of phosphate fertilizers is essential for increasing crop yield. Water solubility of phosphate fertilizers depends on both acidic and neutral to alkaline conditions. Several factors that influence the application of rock phosphate as P fertilizer are rate of dissolution, soil characteristics, plant species and fertilizer.
Factors which determine rock phosphate dissolution rate are its lattice composition, accessory mineral type and particle size. Solubility of apatites increased by substituting CO32− for PO43− in the lattice structure due to decrease a-dimension of the unit cell, and crystal instability [19]. Silverman et al. [20] reported Calcium carbonate as the soluble apatite as compared to other apatites. Rate of its dissolution increases with the concentration of Ca2+ and pH at the surface of apatite, and therefore it reduces the rate of rock phosphate dissolution in soil. The rate of dissolution reduced under field conditions due to leaching or plant uptake of calcium ions. The rate of dissolution increases as the particle size decreases; this might be because fine particle size has greater degree of contact between rock phosphate and soil.
The rate of dissolution of rock phosphate also depends on the chemical properties and type of soil to which it is applied. As compared to other parameters, pH buffering capacity was very important in soil. Earlier studies indicated that the amount of rock phosphate-P decreased with the increase in soil pH. The rate of dissolution of rock phosphate was highly sensitive to Ca2+ activity in the soil solution. A linear relationship between the log of Ca2+ activity and log of P in soil solution has been reported by Robinson and Syers [21]. Phosphate retention capacity and soil moisture also affect rock phosphate dissolution of the soil to retain P. Wet soil enhances the rate of phosphate dissolution by allowing the dissolution products. The product transported away from the surface of the rock phosphate particles and recognized the positive effect of organic matter on rock phosphate dissolution.
Ability of plants to extract P from rock phosphates was recognized by Merril, quoted in Flach et al. [22]. Plants affect the dissolution by the secreting acid or alkali, through Ca uptake, production of chelating organic acids such as citric, malic and 2-ketogluconic acids which complex Ca and deplete P in the soil. Roots of the plants induced change in rhizosphere pH which causes imbalance in the proportion of anionic (usually NO3−, H2PO4−, SO42− and Cl−) and cationic nutrient (K+, Ca2+, Mg2+ and Na+) uptake by the plants. The imbalance in the rhizosphere is maintained by the release of either H+ or OH−/HCO3−, thus balancing the pH of the rhizosphere. Acidic soil enhances the rate of rock phosphate dissolution. Effective rock phosphate utilization by plant species such as e.g., buckwheat and rapeseed has been responsible for their high Ca uptake. Flach et al. [22] assessed the ability of maize, pearl millet and finger millet to utilize P from rock phosphates by a pot experiment and concluded that plant species influence P dissolution; therefore, choice of crop is very important to maximize the solubility of rock phosphate.
Today growing extraction and consumption of phosphate is exhausting existing deposits, and therefore the rate of P reserves depleting. This means that at a certain point time comes when all the phosphorus reached to the alarming peak and that condition is called ‘peak phosphorus’ according to literatures. This condition will be calculated on the basis of phosphate rock reserves. Since no consensus was there on the size of these reserves, so nobody knows when will be the peak phosphorus stage will occur. Peak phosphorus has been calculated by the Global Phosphorus Research Initiative (GPRI). In 2009 the GPRI estimated that phosphorus production would peak around the year 2033 and that afterward production will continuously decrease until reserves are depleted within the next 50-100 years. The US Geological Survey (USGS) re-estimated reserves at 60,000 mmt up from previous estimates of 16,000 mmt; the IFDC stated that ‘there is no indication of “peak phosphorus” event within the next 20–25 years. The concept of peak phosphorus itself is contested; the main fault in the calculations of peak phosphorus is based on phosphate rock reserves not resources which provide the basis for estimation of static ranges. Phosphate rock reserve data explained by national geological surveys do not point out the absolute quantity of an element which is available for extraction, as the static paradigm would suggest. According to the ‘dynamic adaptive paradigm’, due to changes in economic feasibility, scarcity in the production of phosphate rock occurs. This paradigm led to scarcity which is a permanent feature of human existence: minerals become scarce as long as they are immensely valued in the society, and how much time and effort it takes to extract them, and they are related to all other goods and services in the society. Shortage of phosphate rock is an important issue when observed from a different angle that is other than relative availability. One of the reasons of fall in phosphate rock exports is the geopolitical turmoil in supplier regions. Scarcity may also result by lack of water available to the mining industry. Price inelasticity of supply, time and investments are the limiting factors which can lead to scarcity of phosphorous rock.
Nowadays the good news is that crisis can be averted. Almost 4/5 of the phosphorus mined for food production never actually reaches the food on our forks. We can therefore invest in renewable phosphate fertilizers or innovations in on-farm efficiency to safeguard our farmers, our agriculture and food consumers. In every sector recycling of phosphorous is efficiently taking place from agriculture and mining to sanitation sector to changing diet (Figure 2) [23]. To meet long-term phosphorous demand of society, we have to face the technical and institutional challenges for implementing practical solutions. An integrated, context-specific approach should be developed over partial measures. Technologies and practices with effective policy instruments (regulatory, economic, facilitation) are required to encourage and bear such measures.
Sustainable phosphorus measures: Efficiency, recycling and changing diets.
Food demand is rising globally with no slowing down in sight. Especially in China and other rapidly growing economies, more demand for meat and dairy means more demand for fertilizers, while human body only needs around 0.4 kg of P each year. 22.5 kg of phosphate rocks are mined to meet the requirement of phosphate for each person’s diet. For growing population water and energy are considered as critical for meeting future demands of food for increasing population. However, there is no approximated value of phosphorus scarcity in future as a limiting factor. Thus far we can say that without phosphorus, there would be no food and life on earth. There is no single international body responsible for managing global P resources currently in the long term, unlike oil, water and nitrogen.
Phosphorus can be recovered and used over and over again if present in sufficient concentrations dissimilar to oil, which is lost once it is used. Between the phosphate reserves and the food which we eat, up to 80% of P is lost in the process from production of fertilizers, application of phosphate on fields, in food processing and final consumption. With the increasing efficiency of phosphorous, we have to carry on recovery process of P from residues of crops, waste food items in dumpsters, manure, human excreta, struvite and other sources such as bone meal, ash and algae. A key opportunity to meeting the goal of global food security lies in the often overlooked link between addressing hunger and sanitation. In agriculture P plays a critical role as a nutrient, and but on the other hand, it is also considered as an environmental pollutant due to sewage emerging from human settlements. Human activities produce 3 million tons of P each year. If this P is recirculated back in agriculture fields from where they first came, we can maintain balance in sustaining food production in the decoupled communities which are dependent on globalized P fertilizer markets.
It may be concluded that evolution of phosphorous acid (H3PO3) and its salts as fertilizers owes much to both the early investigators searching for phosphate replacements and to the many scientists who later sorted out the relevant facts about plant response to Phi through phytopathological research. In recent years, scientific aroused their interest in the nutrient properties of H3PO3-derived products which stimulate their commercialization of H3PO3 as fertilizers. Trails of these fertilizers on crops have given fruitful results. The nutritive properties of Phi products proved to be a useful addition to producers’ resources. Phi fertilizers elicit positive responses to crop like it enhances flowering and fruit set in cirrus and are converted to phosphate through oxidation process. H3PO3 derivatives give similar responses as that of orthophosphate in fertilization of crops, despite sometimes delayed. Phosphorus acid-derived fertilizers provide a more readily available source of P than that which occurs in soil. Phi products provide more phosphorus to plants as compared to phosphate fertilizers due to their high efficacy of phosphorus uptake through plant foliage. Earlier concept of not using phi as P fertilizers is now changed due to positive results recorded by the scientist about the use of these products as fertilizers. Thus we can reduce a lot of financial burden from our economy by reducing the import of rock phosphate and other by-products required in the manufacture of commercial P fertilizer.
The authors are grateful to section of Ecology and Environmental Botany, Department of Botany, A.M.U, Aligarh.
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