The ABO blood groups of couples used in the experiments
\r\n\t
\r\n\tSince its discovery in the mid-50’s, a wide range of applications from low to high voltage appeared, putting polyimide as a key material to design more performing and reliable electrical devices and systems. On another hand, polyimide appears also essential for the development of new electronic devices where further considerations such as high power density, integration, higher temperature, thermal conduction management, energy storage, reliability or flexibility are required in order to sustain the growing electrical energy consumption needs of the global society.
\r\n\tConsequently, polyimide materials have and will have to face new exciting fundamental, technological and environmental challenges among which:
\r\n\t• a better understanding of its intrinsic electrical properties to identify current limitations and propose new advanced device designs,
\r\n\t• the development of innovative composites and nanocomposites structuration to tailor its physical properties by involving classical and original nanoparticles such as graphene layer, carbon nanotubes, metal, silicates, nitrides, etc.,
\r\n\t• the development of polyimide composites for energy storage, thermal management, reinforced nanodielectrics and corona-resistant nanocomposites,
\r\n\t• the development of new low and ultra-low dielectric constant polyimide for microelectronics (fluorinated polyimides, nanoporous, mesoporous),
\r\n\t• the development of new higher temperature reliable polyimide (high glass transition, high degradation temperature),
\r\n\t• the emergence of solvent-free processes to fit with environmental purposes
\r\n\tMoreover, many challenges regarding the aging mechanisms understanding under single or multiple constraints and the realistic lifetime prediction using robust physical modelling is a ubiquitous questioning in most of the electronic industries.
\r\n\tThis book will target to review both the state-of-the-art and new researches on Polyimide for Electronic and Electrical Engineering Applications. It will present interdisciplinary chapters on the state of knowledge of each topic under consideration through a combination of overviews and original unpublished research. Chapter proposals related to one of the following topics and their keywords (but not only restricted to them) are very welcome to be submitted for this book publication project:
\r\n\t• General Considerations and Technological Processes of Polyimide for Electronics and Electrical Systems
\r\n\tProcessability, Photosensitive and non-photosensitive polyimide, Curing temperature,
\r\n\tSpin-coating, Dip-coating, Extruded enameled wires, Other casting methods
\r\n\t• Polyimide in Microelectronic Applications
\r\n\tDielectric properties, Intermetal layer, Ultra Large-Scale Integration (ULSI), Low-k dielectrics, Fluorinated polyimide, Nanoporous polyimide, Flexible substrates, Thin film transistors, LCD devices, sensors and actuators (gas, humidity, pressure, tactile…)
\r\n\t• Polyimide in Medium and High Voltage Applications
\r\n\tElectrical insulation properties (conduction, breakdown), Digital isolators, Power electronics and devices, Power modules, Power integration, Passivation, Packaging, High voltage power systems, Enameled wires for fed-inverter rotating machine
The only identifiable function of Red Blood Cells (RBC) is the delivery of Oxygen. In mammals, RBC is a unique cell because:
It does not have cellular organelles like any other cells in the body.
It has a very especial protein: Hemoglobin which plays the role of carrying Oxygen to tissues and carries back carbon dioxide to lungs.
It has a very especial cell membrane which carries a number of blood groups antigens’ systems. Their functions include transporting other proteins and molecules into and out of the cell, maintaining cell structure, attaching to other cells and molecules, and participating in chemical reactions [1]. Those systems are genetically controlled with blood groups determining genes.
The work described in this chapter is based on the function carried out by the cell membrane antigens which are transporting other proteins and molecules into and out of the cell. The question is what are those proteins that are transported? In fact, this question identifies the knowledge gap about RBC role in health and disease. In the next section, some hypotheses will be inducted and deduced through analysis of available background knowledge. The experiments that can proof those hypotheses are described in section 3. This is followed by describing a theory about the role of RBC in health and disease based on the proved hypotheses and how we can benefit from this theory in diagnosing and treating of patients.
Basically, when an antigen is introduced into a body, the immune system (IS) does either one of two reactions: immune tolerance (IT) or immune response (IR). IT-reaction is never absolute [2]. It is usually accompanied by a weak IR. In normal IR, one cannot identify if there is a degree of IT, because there is no defined laboratory method/test that can measure the degree of IT. Meanwhile, by logical implication, some degree of IT should exist with the normal IR. This entails that there is an equivalence relation between IT and IR.
Hypothesis I: There is no absolute immune tolerance, if and only if there is no absolute immune response
In central IT, immature self-reactive T lymphocytes recognize antigens in the thymus and undergo negative selection (deletion) [3]. Consequently, in normal IR against a particular antigen, measuring the concentration of this antigen in the thymus can be correlated to the degree of the accompanied IT. The transport mechanism of antigens to the thymus is a critical issue because of the remarkable capacity of IS which can recognize any antigen [4]. In [5], authors claim that Dendritic Cells (DCs) have several functions, not only, in innate and adaptive immunity, but also there is increasing evidence that DCs in situ induce antigen specific unresponsiveness or tolerance in central lymphoid organs and in the periphery. The evidence that DCs transport antigens to thymus in central tolerance is very weak while the evidence that DCs have role in peripheral tolerance is more acceptable based on the review article [6]. In conclusion RBC may be vehicles which transport self antigens to induce central IT.
The role of RBC in transporting antigens has not been investigated before. If RBC are capable of antigen transport to induce IT, this will unveil important knowledge. For instance, in hemolytic disease of fetus and newborn (HDFN), maternal anti D alloantibody and feto-maternal ABO incompatibility are the two major causes of HDFN, Meanwhile, with the implementation of Rhesus D immunoprophylaxis, hemolytic disease due to ABO incompatibility and other alloantibodies have now emerged as major causes of this condition. [7].
In pregnancy, most of delivered infants are normal when there is no anti D alloantibody which means that there is an efficient mechanism that can handle the other incompatibilities. The mechanisms explained in literature explain why ABO incompatibilities, only, do not occur [8], [9] and [10], but these mechanisms do not explain why those incompatibilities occur. The mechanism may be based on trapping those antibodies in placenta through RBC catering of ABO and other incompatible blood groups antigens. Consequently, the occurrence of HDFN may be due to depletion of those antigens’ store from RBC. Also, if this RBC transport function is the mechanism a body tolerates his self antigens, this will explain how a pregnant woman is able to tolerate her fetus and placenta, assuming that they are part of self.
Hypothesis II: RBC hide antigens to transport them to target organs.
From these hypotheses I & II, if RBC play role in antigen transport, one can deduce that in any mammal, blood circulating antibodies against self and foreign, either antigens or tolergens, will react with hemolysate.
Hypothesis III: There is an injection function (one-to-one) between circulating antibodies and RBC\'s hemolysate antigens.\n\t\t\t
To proof that RBC have role in immune reactions (IR and IT), one need to proof that there is an inverse correlation between antibodies concentration in plasma and antigens concentration in RBC.
Hypothesis IV: In immune response, antibodies concentration in plasma against a particular antigen in hemolysate is higher than this antigen concentration in hemolysate. Meanwhile, in immune tolerance, antibodies concentration in plasma against a particular antigen in hemolysate is lower than this antigen concentration in hemolysate.
It should be remarked that Humans expressing a defective form of the transcription factor AIRE (autoimmune regulator) develop multi-organ autoimmune disease (autoimmune polyendocrinopathy syndrome type 1) [11]. Liston et all [12] prove that this autoimmune syndrome is caused by failure of a specialized mechanism for deleting forbidden T cell clones, establishing a central role for this tolerance mechanism.
The methodology applied will demonstrate the existence of particular self tolerogens and particular foreign antigens in RBC (Hypothesis I & II) and show that innumerable antigens exist in RBC which react with innumerable antibodies that exist in plasma. This partially proves that RBC play a role in immune reaction. To proof Hypothesis IV, it will be demonstrated that the concentration of foreign antigens in RBC varies by time in relation to IR known behavior. The experiments done are the following:
RBC of pregnant females transport male spouse ABO blood group antigens
RBC of pregnant females transport male spouse HLA antigens
RBC transport self HLA antigens
RBC transport self Tissue Specific Antigens (TSAs).
RBC hemolysate antigens are precipitated by plasma obtained from the same individual and cross reacted with plasma from different individuals.
RBC transport bacterial antigens.
RBC antigens and plasma antibodies concentration vary with time.
Couples that have children, pregnant females, and single females were selected from relatives and friends. The purpose of the experiments was explained to them. Not all the combinations could be found, after blood grouping. The combinations presented, in Table-1, were used to conduct the experiments. Blood samples were taken on heparin. Some of the blood samples were used to prepare RBC and plasma and the rest was used to prepare lymphocytes using the Ficoll hypaque technique [13].
RBC were washed several times using phosphate buffer saline (PBS). The male RBC were divided into two tubes. The first tube was divided into small aliquots that were frozen to rupture RBC. The second tube was used to prepare a 5% suspension. The female RBC were divided into small aliquots that were frozen to rupture RBC. Notice that we do not need female intact-RBC.
Female ABO group | Male ABO group |
O | A |
O | B |
O | AB |
A | O |
A | B |
B | A |
O (SINGLE) | - |
The ABO blood groups of couples used in the experiments
To test RBC transport of male spouse ABO antigens, a technique based on competitive inhibition of RBC agglutination was followed. If the hemolysate contains ABO specific antigens, then those antigens will compete with RBC and prevent their agglutination. Figure 1 illustrates a schematic description of the experiment.
Method
The experiment was performed, for each couple, as follows:
In positive control tubes which represent also reference tubes for comparison with test tubes, serial dilutions (up to 1/128) of female spouse plasma were made using normal saline. A drop of a male spouse\'s hemolysate was added before adding his RBC’s suspension.
In test tubes, serial dilutions of the female spouse\'s plasma were made using normal saline. A drop of the female spouse\'s hemolysate was added before adding a drop of her male spouse\'s RBC suspension.
Results
Whenever there is ABO incompatibility and the male spouse is not ‘O’, agglutination was inhibited by the female spouse hemolysate and was not inhibited by male spouse hemolysate. In most cases, agglutination was inhibited in the first tube. However, agglutination was never observed in subsequent tubes. The single virgin female RBC do not contain any ABO antigens.
This experiment was performed using commercial HLA Typing Trays for the identification and definition of HLA Class I Antigens using the microlymphocytotoxicity assay [14]. It is also based on competitive inhibition. Consequently, if typing wells that show positive reaction were inhibited in corresponding testing wells by adding hemolysate, this proves the existence of specific competing antigens. Figure 2 illustrate the experiment steps.
Schematic drawing of ABO antigen transport experiment, the upper part shows how the reference positive control is conducted, while the lower part shows how the test is conducted.
Re-typing of male spouse but using his female spouse hemolysate to compete with his lymphocytes
Method
First, each couple was HLA typed, and then the following was done:
A hemolysate from a third person was added to control wells. The positive control should give positive reaction. In this way, we excluded inherent errors or non-specific reaction.
Female spouse hemolysate (diluted 1/16) was added to typing wells
Male spouse lymphocytes was added and followed by the complement and eosin dye as usual.
Wells that gave positive reaction in typing were examined by contrast microscope.
Results
It was observed that female spouse hemolysate inhibited the typing reaction while the third person hemolysate did not. This indicates the existence of male spouse HLA antigens in female spouse hemolysate.
This experiment is similar to the previous one. The only difference is the use of the male’s own hemolysate instead of his female’s spouse hemolysate. It was observed that a male hemolysate inhibited the typing reaction of his lymphocytes indicating the existence of self HLA antigens.
If RBC transport antigens to central organs of the immune system to induce tolerance, then RBC will definitely transport TSAs. Otherwise this transport function has nothing to do with tolerance. Consequently, the objective of this experiment was to demonstrate that antibodies against TSAs can be prepared through injecting RBC of white mice into rabbits. Figure 3 illustrates the experiment.
Preparing antibodies against white mice TSAs in rabbit
Materials
A number of white mice were slaughtered to collect their blood on sodium citrate and their organs (liver, kidney and spleen) were preserved on 10 % formalin. The separated RBC were washed many times with sodium citrate and then diluted with 3% formol-saline to kill any bacterial contamination. An ordinary rabbit was selected to prepare the antibodies.
Method:
A rabbit was injected subcutaneously with one ml of white mice RBC for four times on weekly intervals.
Blood was collected from ear-vein after 35 days from the first injection.
The serum was examined for antibodies against mice RBC using direct agglutination slide test.
The serum was examined for antibodies against TSAs of white mice (liver, kidney and spleen) using the sandwich technique in histo-pathology sections.
Results:
All sections showed florescence. Figure 4 illustrates some of the histopathology sections taken from a white mouse’s organs.
Histopathology sections from a white mouse’s organs examined by florescent microscope showing florescence due to antigen-antibody reaction, A: kidney tissue, B: liver, and C: spleen.
Ouchterlony immuno-precipitation test of normal serum against self and other normal hemolysate was conducted, Figure 5(a). We confirmed this finding by using Western Blot technique, and showed that serum from one individual recognized antigens in hemolysate from two normal persons, Figure 5(b). Further confirmation was obtained by using two-dimensional gel electrophoresis (2-DE) of co-immunoprecipitated hemolysate antigens using self-serum, Figure 5(c). Notice that the number of the immune-precipitated antigens is numerous and many spots were enriched by immune-precipitation because those antigens were not detected in 2-DE gel of hemolysate, Figure 5(d). Antigenicity of the separated proteins was confirmed by immune-blotting proteins separated by 2-DE with the same self-serum, Figure 5(e). This excluded co-precipitation of non-antigens, as they would not be detected in immune-blotting.
As TB is a priority disease, trying to find Mycobacterium tuberculosis bacilli protein antigens (MTPAs) in TB-patient hemolysate was conducted through 2D electrophoresis, and then identifying gel spots with mass spectrometry. Fortunately, we discovered four MTPAs. This motivated us to do the experiments of the next section to identify more MTPAs in hemolysate of TB patients.
Identifying MTPAs in TB patients hemolysate
The goal is to find the set of antigens, in TB patients’ hemolysate, which is related to Mycobacterium tuberculosis bacilli. The approach taken follows the following steps Figure 6:
The study resources are:
[A] Patients
[B] Mycobacterium tuberculosis (H37Rv)
For each patient:
Collect blood sample on anticoagulant (step 1)
Separate RBC and wash many times with saline (step 2)
Hemolysate [C] is prepared by rupturing RBC with low isotonic solution which is the binding buffer in affinity chromatography
Prepare hyper immune serum for M. tuberculosis (step 3)
Purify antibodies using Protein A Sepharose beads (step 4)
The purified antibodies are then used to separate antigens from hemolysate (step 5)
The disease related antigens are identified using in gel trypsin digestion and MALDI TOF mass spectrometry (step 6)
Detection of innumerable antigens in Red Blood Cells. (a) Ouchterlony test showing serum of normal against hemolysate of self and others (b) Western Blot using two normal hemolysate propped with serum of one of them (c) Silver stain of 2-DE of immune-precipitated hemolysate antigens (d) Silver stain of 2D electrophoresis of hemolysate (e) Western Blot of 2-DE of hemolysate propped with serum.
The objective of this experiment is to investigate the dynamics of foreign antigens in RBC. In effect, antibodies are taken at one instance of time, while RBC are taken at different instances.
Flowchart depicting the resources and steps for identification of hemolysate antigens related to Mycobacterium tuberculosis (H37Rv)
Precipitated Antigens separated using 2D electrophoresis Gel
Materials
Bacteria: Escherichia coli O157:H7 strain[1] - was inoculated onto SMAC agar (Oxoid). Colonies were tested by E. coli O157 latex kit (Oxoid DR 620) and confirmed biochemically. A single colony of E. coli bacterial growth from the plate was inoculated into Brain heart infusion broth (Oxoid) and incubated overnight at 37 ˚C and adjusted to a concentration of approximately 1010 CFU.
Animals:
A rabbit
A baladi sheep between 8 to10 months.
The rabbit and sheep were tested serologically, to be negative, for Escherichia coli O157:H7.
Methods:
The first experiment method was done as follows, Figure 8:
Rabbits were vaccinated by Escherichia coli. Rabbits were injected subcutaneously with one ml on weekly basis for three weeks.
Blood was collected from the ear-vein after 21 days from the first injection.
Rabbits sera were separated and examined for antibodies against E. coli O157 using direct bacteria slide agglutination test.
A sheep was infected by oral administration of bacterial suspension.
Red blood cells were prepared from anti-coagulated sheep blood collected at 0 time (i.e., before inoculation), 1st week, 2nd week, and 3rd week. The collected blood was centrifuged at 4 ˚C for 25 minutes at 1170 g. Plasma and Buffy coat from each sample were removed. RBC were washed twice in normal saline solution by centrifugation at 4 ˚C for 5 minutes at 2000 g, and then re-suspended in Tris/Saline buffer pH 7.5 and subjected to lyses by freezing.
Nobel agar 1% in Tris/Saline was used as a supportive media for antigen-antibody precipitation, where the central well contained rabbit serum and peripheral wells contained sheep RBC hemolysate.
Results
The rabbit serum showed high titer (1/160) of antibodies against E. coli. Antigens of E. coli could be precipitated from sheep RBC of the 1st and 2nd week after infection, only, Figure 9.
Preparing Antibodies against bacteria and preparing RBC carrying antigens of this bacteria. The purpose is to precipitate Bacteria antigens from RBC of infected animals using the prepared antibodies against those antigens.
Illustrates the dynamics of RBC’s antigens
The RBC transport function maintains tolerance to self antigens. This function is exploited positively to protect a fetus from the immune system attack using the same mechanism of protecting the self. In effect, a fetus, which is an allograft, is considered part of self.
In humans and animals, not all microorganisms are capable of causing disease. Some of those microorganisms are equipped with the machinery that can overcome biological barriers and can cause disease in animals but not in humans and vice versa [15]. The role of RBC antigens transport in inducing tolerance to self-antigens is a feature that can be considered as a security-hole, as invaders can exploit this process to escape from the response of the immune system by disguising themselves as self. Tumors and parasites are negative examples.
Notice that this mechanism of tolerance induction does not contradict with all what we know about tolerance. Further, it explains the documented properties of tolerance. For instance, some of the properties that can be explained are:
Artificially induced tolerance is of finite duration because antigen stores get depleted.
Tolerance to self antigens is a process that continues throughout life but begins during fetal development because RBC are transporting self antigens all the time.
Notice that the discovered function of RBC fills a gap in the understanding of tolerance. Part of this gap can be expressed in the following questions:
Why soluble antigens administered intravenously favor tolerance while particulate antigens injected into the skin favor immunity.
Why ingested large doses of soluble proteins induce systemic T lymphocyte tolerance, whereas the components of vaccines such as the Sabin polio vaccine induce an effective local immune response.
Why tolerance is easier to induce in prenatal rather than postnatal life.
Answer of Question 1 and 2: RBC can easily absorb soluble antigens through pinocytosis while a particulate antigen needs receptor sites on RBC in order to be absorbed, which is the RBC membrane antigens function. Notice that the probability that the immune system will react to some processed antigens still exists. That is why the dose of antigens plays an important role. As far as there are enough stores of antigens in RBC, they are effectively tolerated.
Answer of Question 3: If antigens are introduced to a fetus while the immune system is still incapable of respond, there is a good chance for those antigens to be processed by the Antigen-Presenting-Cells (APC) and then absorbed by RBC. When mature lymphocytes production starts, later in life, antigen stores of RBC are used to induce tolerance. This may explain why tolerance is easier to induce in prenatal life.
Further, a pathogenesis mechanism of some autoimmune disease can be postulated. If RBC antigen-transport function is impaired for a particular self-antigen, for some reason, the tolerance to that antigen will eventually vanish. Consequently, an autoimmune response will be provoked to that antigen and autoimmune disease is established.
This observed RBC antigens transport function creates an antigens’ store. This store can be exploited in many directions. The proposed direction is to exploit functional proteomics approach [16] with the following three crucial aspects of the experimental design to produce products which are among diagnostic kits, vaccines or treatment components:
The strategy used for the selection, purification and preparation of the antigens to be analyzed by mass spectrometry
The type of mass spectrometer used and the type of data to be obtained from it
The method used for the interpretation of the mass spectrometry data and the search engine used for the identification of the proteins in the different types of sequence data banks available
The aim of this approach is to identify antigens which are relevant to a particular disorder.
This approach is based on using a subset of antibodies which are specific against a subset of antigens of a particular disease to enable the use of those antibodies and those antigens in preparation of beneficial products.
Diagnostic kits can be prepared for all infectious microorganism and all tumors. In such disorders, simple kits can be prepared using the following steps:
Extract antigens from microorganism/tumor-cell-line cultures in coupling buffer
Prepare hyper immune serum using extracted antigens
Build an affinity column
Antibodies purification: Use affinity column containing antigens to separate their related antibodies from hyper immune serum
Adsorb purified antibodies to latex beads
A more advanced kits based on selection of antigen-determinant sites (epitopes) can be prepared. The problem of such kits, which uses a particular antigen, is in its validation which will be more sophisticated. One can expect that this particular antigen may not exist in RBC antigens’ store of some population who are genetically different from the population used in preparation of the kit.
Active vaccines against all infectious microorganism and all tumors can be prepared by using the purified antibodies prepared for diagnostic kits in identifying related antigens existing in RBC antigens’ store. The identified antigens can be prepared using the technology of recombinant proteins purification.
The proposed mathematical model and a data mining algorithm will not only help in identifying proteins (antigens) that can be used in diagnosis and treatment of difficult disorders, but also will help in etiological diagnosis of idiopathic disorders and their treatment. This approach is based on building large databases of RBC antigens’ store for patients and normal individuals. Consequently, a patient sample is collected on anticoagulant. RBC and plasma are separated. The plasma IgG is separated and then used as ligand in immunoaffinity chromatography to separate hemolysate antigens. The collected antigens are identified by mass spectrometry. The database record consists of the diagnosis and the set of identified antigens.
It consists of four main parts; definitions of symbols, model of diseases caused by microorganisms, tumors, or foreign proteins; model of autoimmune diseases which result as a consequence of missed tissue proteins from RBC antigens’ store; and model of diseases of unknown cause (Idiopathic).
Definitions
Let the assumption of this work be as the following:
pi: protein amino acid sequence, where i = 1.. n
dj: health state, i.e., normal or disease name, where j = 1.. m
P = {p1, …, pn}, Set of all proteins of RBC antigens’ store
D = {d1, …, dm}, Set of all diseases
Pp: patient proteins where Pp P where p is the patient ID
Op: (pi, dj), ordered pair of patient presented by protein sequence (i) and health state (j).
Model of Diseases caused by microorganisms, tumors, or foreign proteins Pdj = ∩{Pp}dj
Where Pdj is the set which contains all common proteins associated with dj. Pnormal = {Pp}normal\n\t\t\t\t\t
Where Pnormal is the set which contains proteins associated with normal.
P\'normal such that p in Pnormal if the number of occurrence of p Pnormal is less than 5% of the total number of p in Pnormal then remove p from Pnormal.P\'dj = Pdj – P\'normal
Where P\'dj is the set which contains proteins that can be used as biomarker or vaccines, Figure 10.
Venn diagram depicting set of abnormal protein of disease X (P\'dj)
Model of Diseases caused as a result of missed tissue proteins
Pudj =
The result is the set which contains all proteins associated with dj
P\'\'dj = Pnormal - Pudj
The result is the set which contains proteins that can be used to diagnose patients through detecting circulating auto-antibodies and to treat those patients through desensitizing them with the proteins that give positive reaction, Figure 11.
Venn diagram depicting set of missed normal proteins of disease Y (P\'\'dj)
Model of Diseases of unknown cause (Idiopathic)
There are many diseases that are identified as idiopathic.
Those diseases can be caused due to existence of abnormal protein or absence of tissue proteins.
Applying data mining methods (A and B) can help to identify new diseases and treat patients appropriately.
Patients’ blood samples will be collected on anticoagulant. RBC and plasma are then separated in different tubes. Plasma is used as a ligand in immune-affinity chromatography to separate hemolysate antigens that can bind to plasma antibodies. The separated antigens are identified by MS and stored in the database indexed by the patient disorder.
In the same time, queries are done to verify the diagnosis and get a prognosis and a recommended treatment component. The following formulas describe the usage of this model in clinical practice.
Let Dp\' is the set of all discovered P\'dj
Let Dp\'\' is the set of all discovered P\'\'dj
Then
Else
P\'\'dj Dp\'\', if P\'\'dj\n\t\t\t\t\t\t
All the previous work in RBC proteomics neither has identified another function nor has mentioned the finding of: HLA, TSAs, or foreign proteins. The reasons are obvious. Firstly, it is not expected to find such proteins and consequently the method used for the interpretation of the mass spectrometry data, and the search engines used for the identification do not consider the right types of sequence data banks available. Secondly, the amount of most of the antigens which belong to the RBC antigens’ store is little. This makes those antigens invisible and hence easily missed.
The work described is just a pilot study that throws some light on a new theory related to RBC. This theory is based on finding antigens’ store consisting of self and non-self antigens. Although this theory can be related to immune tolerance by logical induction, the concrete evidence and mechanism need further research. Mainly, the logical induction is based on finding all kind of antigens in hemolysate, especially HLA antigens which are related to fetus. This existence of all kinds of antigens, definitely, plays some immunological role which may be immune tolerance.
The initial experiment, which shows the existence of ABO antigens in hemolysate of pregnant females, explains the mechanism of how HDFN occurs. Meanwhile, the experiment which shows that HLA antigens exist in their hemolysates proposes a new mechanism by which a pregnant woman is able to tolerate her fetus and placenta. Simply, it is the same mechanism a body tolerates his self antigens.
The experiments which use hemolysate against self-serum: Ouchterlony immune-precipitation test, Western Blot, and 2-DE of co-immunoprecipitated antigens demonstrated that RBC have an antigens’ store. Mass spectrometry of spots obtained from 2-DE gel demonstrated the finding of all kind of antigens, self and non-self, in hemolysate. This indicates that blood circulating antibodies in any individual will react with his RBC’s hemolysate antigens. In effect, there is no absolute immune response, too.
This directed our attention to use hyper immune serum against Mycobacterium antigens. This will help to get rid of other proteins and do better separation; and hence better identification. Consequently, we could identify 11 proteins from 60 gel spots belonging to H37Rv strain. The rest of spots are proteins related to bacterial commensals. Consequently, purification of specific antibodies from hyper immune serum is recommended to get further better separation.
In the experiment which investigates the dynamics of foreign antigens in RBC using sheep RBC which has been infected with E. coli, it was shown that the concentration of foreign antigens in RBC varies by time in relation to IR known behavior. This proves that RBC have role in immune reactions (IR and IT).
Whatever the reason of this existence of antigens in hemolysate this existence can help in designing diagnostic kits for different types of diseases. Further, it will help in discovering, not only, new immunological disorders which are, now, categorized under idiopathic disease, but also, identifying the obscure cause of many immunological disorders, including cancer. The identification of the cause of a disorder will help in its treatment and prevention.
There are some major transformations taking place in mature tourist destinations. These destinations correspond to the places with large tradition on touristic activities and situate in the mature or decline phases of the cycle of life of touristic destination [1] to refer the sun and beach tourism as Tenerife Island. These are the results of both the obsolete nature of tourism services, equipment, and infrastructures, as well as changes in the profile of demand for more sustainable products. Both transformations have motivated tourism agents to seek innovative, creative, and imaginative solutions that prevent the decline of destinations and that represent a claim for visitors. The main examples are the changes in the touristic laws associate to the sustainability concept, the transformation of the urban spaces in the private and public places in the touristic cities, and the creation of the new touristic products such as geotourism. In this sense, within these new motivations, the diversification of the offer in the destination is being key. Therefore, the creation of new tourism products and experiences has become one of the main challenges for mature destinations to continue to occupy a significant role in the world tourism map at a time when the tourism sector is being very dynamic.
Within the diversity of new products and tourist experiences that are being developed in many parts of the world [2, 3], is geotourism. Although it is true that in its landscape and esthetic conception, relief was present in the declaration of the first national parks in the world such as Yellowstone, Grand Canyon, etc., and in Spain (Picos de Europa, Teide, etc.), it is not until recent times geotourism has increased significantly [4, 5, 6, 7], contributing to the creation of the European and global networks of geoparks in the last 20 years. Therefore, geotourism is a relatively modern term that was implemented in the modern society but welcomes multiple initiatives in different places and from different geological and geographical approaches, but they do not have to be mutually exclusive [7, 8].
The island of Tenerife is an international tourist destination whose main attraction is the sun and the beach that attracts millions of visitors every year. However, Tenerife has a great variety of natural resources and tourist attractions that could diversify its offer and be the claim of other types of tourists [9]. In this sense, this chapter shows the main results of the project “Tfgeotourism: strengthening the economic and business fabric linked to the tourism sector of Tenerife by promoting volcanotourism,” whose objective is to create different geotourism products and experiences that contribute to diversifying the offer of leisure activities on the island and that can be implemented by tourism companies that already exist in Tenerife or that may be created as a result of this offer of volcano tourism products.
Therefore, the general objective and the main results of this project are in accordance with the objectives of the Global Geopark Network (GGN) to develop a leisure offer supported by geotourism that contributes to diversifying tourism and traditional economic activities in Tenerife through the creation of innovative local companies, new jobs, and highly qualified training courses for the different tourism agents [10]. But at the same time, Tfgeotourism is also in line with some of the objectives proposed by the Geoparks Commission of the International Geographical Union (IGU), which relates them to their role as a tool to communicate, recreate, and conserve nature or to understand the wide range of processes that affect the sustainable development of geoparks, including the natural environment, political, and socioeconomic processes [11].
In this sense, the choice of volcanic geotourism within the diversity of new tourism products and experiences in Tenerife responds to several reasons: 1. The island is a consolidated and well-known international destination to which several million tourists arrive annually. 2. The volcanic landscape of the island is diverse and, in it, practically all the forms and eruptive processes can be recognized, giving rise to one of the most geodiverse volcanic spaces from the point of view of the geological and geomorphological heritage [12]. 3. Tenerife already has a geotourism offer in some places such as the Teide National Park, being the most visited park in the Canary Islands and Spain, with an average of about 3 million annually [13, 14], and also in other places such as Barranco de Masca, the Macizo de Anaga, or the volcanic tube of La Cueva del Viento. 4. A large part of the insular volcanic geoheritage is not being exploited for tourism, because the principal activities associate to the geotourism focused in the national park and some natural protected areas. 5. The practice of geotourism does not necessarily require a specialized public, which makes it very attractive and feasible for all tourists and visitors who come to Tenerife.
Tenerife is the largest (2034 km2) and highest (3718 m a.s.l.) island of the Canary archipelago (Figure 1). It has been built up as a result of the accumulation of a wide variety (mafic, felsic) of fundamental volcanic materials in a relatively short period of time, which results in a world-unique variety of volcano-related tourism features in an accessible scale. In Tenerife, this great geodiversity includes the stravolcanoes, shield volcanoes, calderas, cinder cones, maars, tuff cones and rings, and lava fields, all exposed beautifully in cliffs, ravines, beaches, deposits, etc. (Figure 2). This volcanic geodiversity is a nonrenewable heritage [15] with various main characteristics associated with its natural and cultural heritage [16, 17]. The diversity of forms and processes of relief both directly related to volcanism as well as erosion and accumulation make up the volcanic heritage [18] and are responsible for the geodiversity of volcanic geoheritage of Tenerife Island.
Location of Tenerife volcanic island. Source: Own elaboration from the DEM base of Grafcan.
Different volcanic landscapes of Tenerife. (1) Teide and Pico Viejo stratovolcanoes. (2) Las Cañadas caldera and Majua dome. (3) Arafo historical eruption. (4) cinder cone of Montaña taco. (5) last eruption of Tenerife: Chinyero cinder cones (1909). (6) Montaña Amarilla tuff cone and Bandas del Sur pumice formation. (7) Montaña Pelada tuff ring and monogenetic volcanic field. (8) Punta de Teno lava delta. (9) ravines in Teno shield volcanism.
Tenerife is one of the main tourist destinations in the world in relation to the number of accommodation places (>137,000 in September 2019) and the number of visitors it receives, which in 2019 exceeded 5 million [19]. We must also add the local visitors and hikers who increase this number. In turn, Tenerife has a great variety of new tourism products and experiences (geotourism, astrotourism, gastronomic, sports, health, bird watching, sailor, hiking, diving, whale watching, etc.), which are associated with diversity of attractions that the island has. Therefore, Tenerife has a diversified tourist offer capable of responding to this increasingly informed, demanding, and active demand, which makes the island an ideal destination for the introduction of new tourism products and experiences such as geotourism.
Tfgeotourism Project started in 2017, thanks to the financing of the Island Council of Tenerife. The entity responsible for executing it is the Instituto Volcanológico de Canarias (INVOLCAN). INVOLCAN, an entity unanimously demanded by the Senate (11/02/2005), the Parliament of the Canary Islands (01/11/2006), and the Congress of Deputies (12/02/2009), aims to contribute to the improvement of volcanic risk management in Spain (say the Canary Islands; the only volcanically active region of the national territory of Spain with volcanic risk) and the optimization of the management of the many benefits that come with living in a volcanic territory (geothermal resources, geotourism, etc.). Its mission and vision are intended to contribute to the sustainable development of the Canary Islands as well as other volcanic regions.
Within the TFgeotourism project and in accordance with its objectives, different products have been created associated with volcano tourism. This type of tourism in Tenerife constitutes a relatively new modality that consists of the tourist exploitation of natural and cultural aspects directly or indirectly linked to volcanoes. In this sense, the geotourism interest of volcanoes is associated with three fundamental aspects: (1) the landscapes they generate where the esthetic beauty of the processes and forms cannot be compared to other geological territories; (2) the cultural aspect of volcanoes in relation to the societies that agree with them (archeology, history, music, folklore, gastronomy, etc.), configuring volcanic landscapes as key elements and icons in the culture of the peoples that coexist with them and its surroundings; and (3) the possibility of experiencing the power of seeing an erupting volcano as long as the safety of visitors is guaranteed.
Associated both with the objective of the project and with the geotourism interest of the volcanoes, TFgeotourism has created various products and tourist experiences that diversify both the leisure offer of Tenerife as well as geotourism. Each of the products generated within the project and whose purpose is to strengthen the economic and business fabric of the island are listed and characterized below.
The need to make a documentary exclusively about the volcanic geoheritage of Teide volcano responds both to its international importance and to the fact that it had not been done until now. The global relevance of this volcano is due to the following facts: it is the National Park that receives more tourists in Spain (Figure 3) and is one of the most visited volcanic areas in the world [20]; it was part of the 16 volcanoes of the decade in 1990 according to the International Association of Volcanology and Chemistry of the Earth’s Interior (IAVCEI) for the reduction of natural disasters; and it was one of the five European volcanoes (European Laboratory Volcanoes: Teide) of the European Commission, and since 2007, it has been a UNESCO World Heritage Site. To all these recognitions that value the volcanic heritage of the national park through its rich geoheritage, we must also add its varied and unique biodiversity and its rich cultural heritage from pre-Hispanic times to the present day [21].
Visitors to Teide National Park. Source: ISTAC; Istac: http://www.gobiernodecanarias.org/istac/. Own elaboration.
Therefore, despite the relevance of its geoheritage and its tourist importance (international, national, and local), the national park did not have a specific and easily acquired tourist product for visitors who are attracted by the geotourism of Teide volcano. In this way, the documentary (Figure 4) that has been made is entitled “Teide, the sleeping giant”; it lasts for about 30 min, and throughout it, each and every one of the natural elements and cultural that make up the geodiversity of volcanic geoheritage of Teide volcano are displayed: stratovolcanoes, domes, cinder cones, hornitos, craters, lava flows, lava tubes, ravines, periglacial forms, and processes.
Some photographs about the shooting of the Teide documentary with drones. All pictures show the Teide and Pico Viejo stratovolcanoes and Roques Blancos lava dome. Source: Alas.
The documentary was presented in 2019 at the twenty-first edition of the Canary Islands International Medium-Environmental Film Festival (FIMEC) in Garachico, in the north of Tenerife, with a large influx of audiences. From there, it has been screened in various municipalities on the island (La Orotava, Guía de Isora, Los Realejos, etc.) and the objective is that it can be screened in all of them.
This is Tenerife’s first geotourism guide (Figure 5). The main objective is to show the diversity of the volcanic geoheritage of Tenerife through the selection of various places that collect aspects of its geodiversity, its geoheritage, and its geoconservation. According with the more geographical vision of geotourism, the places listed in the guide show the values of the natural and cultural heritage associated with volcanoes. In this sense, and in addition to the geological and geomorphological heritage, aspects related to the vegetal landscape, the ornithological importance, the panoramic views or the cultural legacy that anthropic uses have left on the landscape are pointed out when using materials from the gea for the farming, the stone roads or the traditional buildings.
Tenerife geotourism guide. Source: http://tfgeoturismo.involcan.org/.
The guide has a selection of 50 places of geotourism interest Tenerife [22] (Figure 6) corresponding to the geological context of buildings and volcanic morphologies of the Canary Islands defined according to the Geological Survey of Spain (IGME). With the purpose of a greater understanding, the information of each place is presented in a file format. This has several sections in which the name of the geotourism place of interest and its UTM coordinates appear; the geological context and its main interest (tectonic, morphological, stratigraphic, and petrological); the municipality to which it belongs and its numbering; the location map; its description; the conservation and uses in it in order to establish what can and cannot be done; the trails that run through each place; and the selection of the main geotourism interests of the site and the bibliographic references where you can find more information about each of the places included in the guide.
Spatial distribution of the 50 sites of geotourism interest. Source: http://tfgeoturismo.involcan.org/.
The set of 50 geotourism sites selected from the guide represent the geodiversity and volcanic geoheritage of the island of Tenerife from the topographic, volcanic, morphological, and landscape point of view [22, 23]. Topographically, the two slopes (north and south) and the different altitude levels of the island are represented. Volcanologically, the 50 places are distributed by all the volcanic buildings of Tenerife (shield volcanism-ancient volcanic massifs, rift volcanism and the central complex Cañadas-Pico Viejo-Teide), they represent both magmatic and hydromagmatic volcanism and the different dynamics and behavior eruptive (effusive, explosive and mixed). Morphologically, the places are part of both the forms and processes of direct volcanism (eruptions, volcano fields, lava fields, stratovolcanoes, etc.) and the processes of erosion and accumulation (ravines, cliffs, beaches, dunes, slopes, etc.). And, finally, from the landscape point of view, an attempt has been made that places of geotourism interest are present in all the large landscape units defined for Tenerife, which are the result of the combination of topographic, volcanic, biogeographic, and cultural criteria (landscapes of coastline with beaches and/or cliffs with xeric vegetation, embedded and open ravines with laurel and pine forests, badlands and lajiales without vegetation, volcanoes and lava with pine forests, stratovolcanoes of peaks with high mountain scrub, etc., and on which you cannot ignore all their cultural baggage) such as farming, traditional villages, churches, etc. In order to make the selected places accessible, it has also been tried that they are always within the usual itineraries (routes and circuits) of visitors to the island [23].
In most cases, geotourism products, experiences, and activities are designed to be carried out mainly in protected natural areas. However, geotourism initiatives within cities are gaining more and more interest [24, 25, 26]. In this sense, the urban geotourism is a type of tourism which concentrates on the exploitation of the attractive parts of the relief of the cities. To do so, it is necessary to identify the places, areas, and/or elements with a geological-geomorphological interest (geosites or geomorphosites) and then design geotouristic itineraries which can be of chronological, geological, or spatial interest. The geographical relief can be found in any particular location within the city, either natural (volcanoes, lava flows, ravines, cliffs, beaches, etc.), which means those that have not been occupied by urban constructions, or those belonging to the cultural heritage (roads, buildings, etc.). The urban geotourism is recommended to a very diverse public, from local residents to visitors attracted by the geological heritage.
Within the TFgeotourism Project, 23 urban geotourism itineraries were designed in the different municipalities of Tenerife (Figure 7). In each of them, the natural geotourism sites and cultures of the cities and their environments were identified, selected, inventoried, and characterized [27]. The information of each site was collected in a file whose objective is to provide more information on the places selected to make the route. The itineraries are always developed through the cities, they are easily accessible, with little unevenness, they can be developed at any time of the year given the subtropical climate of the island, the duration varies between 1.5 and 3 h and the number of places and stops is not more than 16.
Example of an urban geotourism itinerary. Source: http://tfgeoturismo.involcan.org/.
On June 19, 1799, the German naturalist Alexander von Humboldt made a stop in Tenerife with the aim of ascending the Teide volcano [28]. The traveler made various observations, descriptions, and measurements of the places he visited. Among the varied information that he left us, stand out his impressions of the natural and cultural heritage associated with the volcanic landscapes through which the route ran from Puerto de la Cruz to the top of Teide stand out. In this sense, his interesting observations have allowed us to identify the 18 stops into which the journey was divided and to show the tourist interest of each one of them. In addition, given the importance of travel literature for experience tourism and the geotouristic interest of the stops on the route (Figure 8), it has been possible to recreate almost exactly the itinerary that Alexander von Humboldt made through the Orotava Valley and the Cañadas del Teide.
Special identification of Humboldt stops on his ascent to Teide. Source: Own elaboration from the DEM base of Grafcan.
Table 1 shows the 18 stops identified and their geotourism attractiveness divided between natural and cultural heritage. The attractions refer both to the impressions taken by the German traveler in June 1799 and to those that can be observed today when the tour is made [29].
N° | Stop | Geotouristic attractions |
---|---|---|
1 | Puerto de La Orotava (Puerto de la Cruz) | Coast of La Paz, hornitos and scorias cones at Tigaiga eruption, Teide, clouds sea, taste for nature, lyrics, music, and San Juan festival in Sitio Liter |
2 | Casa familia Cólogan (Hotel Marquesa) | Basaltic volcanic rocks: lavas, lapilli, etc. |
3 | Jardín Aclimatación (Jardín Botánico) | Topographic terrace, plants, natural source, climate |
4 | Volcanes Fraile y Horca | Geology and geomorphology, volcanic cones, craters, lavas, volcanic ash, lapilli, pumice, basalts, eruptions, vegetation, guanches, and rural landscapes |
5 | Villa de La Orotava | Topography, geology, hills-watersheds, climate, mist, waters, vegetation, dragon tree, ferns, fauna, guanches, noble houses, aqueducts, mills, crops, vineyards, fruit trees, and peasants |
6 | Monte de los Castaños | Cobbled path, chestnut trees, laurel forest, heather, and ferns |
7 | Pino Dornajito | Panoramas of the north of Tenerife, rugged topography, mountainous area, volcanic mountains, volcanic cones, lava, small ravines, porous rocks, water spring, vegetation, pine trees, and footpath |
8 | Monteverde | Heather, fern, fern-rooted gofio, humble town, and misery |
9 | Portillo | Volcanoes, basalts, volcanic cones, caldera, and pine forests |
10 | Cañadas Occidentales-Volcán M. Corrales | Topographic plain, plateau, pumice fields, sands, Teide, lava flows, eruptions, broom thickets, summit vegetation, hunters, and straw hats |
11 | Roque de Gaita | Volcanoes, black streams, obsidian blocks, rocks, rabbits, and goats |
12 | Estancia de los Ingleses-Montón del trigo | Black streams, ravines, plateaus, lava blocks, caves, Teide, climate, gorse, horses, mules, bonfires, and pine torches |
13 | Altavista | Topography, slopes, escarpments, lava, snowfields, snow, mules, and indigenous |
14 | Malpaís (cueva del Hielo) | Malpaís, lavas, path, perpetual snow, glacier, cave, volcanic tubes, jameos, sea of clouds, and ice collection |
15 | Senda coladas negras | Black lava flows, aa, malpaís, python, sea of clouds, vision of other islands, sunrise, laziness of the guides, and guanches |
16 | Rambleta | Topographic plain, python, Narices del Teide, and fumes-gases |
17 | Pitón | Topography, slopes, lavas, python, lavas, cooling sidewall, ash, pumice, slag, snow, and groundwater |
18 | Cráter terminal | Small crater, lava flows, slags, ashes, slopes, sensation of cold, roaring fumaroles, Teide violet, climate, sky, bees, feeling of loneliness, and Malvasia wine |
Identification of the attractions with geotourism interest in the stops made by Humboldt from the port of La Orotava to Teide.
With the aim that all the products produced within the TFgeotourism project are freely consulted and available to everyone, a web page was created (
This chapter shows the different volcano tourism products created within the TFgeotourism project for the island of Tenerife. This Spanish island is one of the main tourist destinations in the world for both the number of tourist places it offers and the millions of tourists it receives each year. The main tourist attraction on the island is the sun and the beach. However, in Tenerife, there is a great variety of tourist resources that could diversify its leisure offer, helping to avoid or minimize some of the problems that mature sun and beach tourist destinations are having. Within the variety of new tourism products and experiences that can be recognized in Tenerife, geotourism associated with volcanoes is one of them. The TFgeotourismo project aims to create different geotourism products to strengthen the economic and business fabric of the island. In this sense, this chapter indicates and characterizes the main activities of the project associated with the documentation and quantification of the conservation values of the volcanic heritage, as follows: (1) production of a documentary on the volcanic geoheritage of Teide volcano; (2) identification, inventory and selection of the top 50 sites of geotouristic interest on the island associate to geomorphosites; (3) creation of urban geotourism itineraries in the different municipalities of the island of Tenerife; (4) recreation of the itinerary of the pioneering German naturalist, Alexander von Humboldt, from Puerto de La Cruz to the top of the Teide volcano; and (5) creation of a web page for the project (
This work is part of the research project “Strengthening the economic and business fabric linked to the tourism sector of Tenerife through the promotion of volcano tourism,” financed by the Program Tenerife Innova of the Cabildo Insular de Tenerife. We appreciate the comments and suggestions of the editor and the two anonymous reviewers. The authors thank David Calvo Fernández for his work throughout the development of the project.
The authors declare no conflict of interest.
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