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Red blood cell
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Red blood cell
Scanning electron micrograph of human red blood cells (ca. 6–8 μm in diameter)
Details
Function
Oxygen transport
Identifiers
Acronym(s)
RBC
MeSH
D004912
Anatomical terms of microanatomy
[edit on Wikidata]
Red blood cells (RBCs), also called erythrocytes, are the most common type of blood cell and the vertebrate's principal means of delivering oxygen (O2) to the body tissues—via blood flow through the circulatory system.[1] RBCs take up oxygen in the lungs, or gills of fish, and release it into tissues while squeezing through the body's capillaries.
The cytoplasm of erythrocytes is rich in hemoglobin, an iron-containing biomolecule that can bind oxygen and is responsible for the red color of the cells. The cell membrane is composed of proteins and lipids, and this structure provides properties essential for physiological cell function such as deformability and stability while traversing the circulatory system and specifically the capillary network.
In humans, mature red blood cells are flexible and oval biconcave disks. They lack a cell nucleus and most organelles, in order to accommodate maximum space for hemoglobin; they can be viewed as sacks of hemoglobin, with a plasma membrane as the sack. Approximately 2.4 million new erythrocytes are produced per second in human adults.[2] The cells develop in the bone marrow and circulate for about 100–120 days in the body before their components are recycled by macrophages. Each circulation takes about 60 seconds (one minute).[3] Approximately a quarter of the cells in the human body are red blood cells.[4][5] Nearly half of the blood's volume (40% to 45%) is red blood cells.
Red blood cells are also known as RBCs, red cells,[6] red blood corpuscles, haematids, erythroid cells or erythrocytes (from Greek erythros for "red" and kytos for "hollow vessel", with -cyte translated as "cell" in modern usage). Packed red blood cells (pRBC) are red blood cells that have been donated, processed, and stored in a blood bank for blood transfusion.
Contents
[hide]
1 Structure
1.1 Vertebrates
1.2 Mammals
1.3 Human
2 Microstructure
2.1 Nucleus
2.2 Membrane composition
2.2.1 Membrane lipids
2.2.2 Membrane proteins
2.3 Surface electrostatic potential
3 Function
3.1 Secondary functions
3.2 Cellular processes
4 Life cycle
4.1 Creation
4.2 Functional lifetime
4.3 Senescence
5 Clinical significance
5.1 Disease
5.2 Transfusion
5.3 Tests
5.4 Separation and blood doping
6 History
7 See also
8 References
9 External links
Structure
Vertebrates
There is an immense size variation in vertebrate red blood cells, as well as a correlation between cell and nucleus size. Mammalian red blood cells, which do not contain nuclei, are considerably smaller than those of most other vertebrates.[7]
Mature red blood cells of birds have a nucleus, however in the blood of adult females of penguin Pygoscelis papua enucleated red blood cells (B) have been observed, but with very low frequency.
Almost all vertebrates, including all mammals and humans, have red blood cells. Red blood cells are cells present in blood in order to transport oxygen. The only known vertebrates without red blood cells are the crocodile icefish (family Channichthyidae); they live in very oxygen-rich cold water and transport oxygen freely dissolved in their blood.[8] While they no longer use hemoglobin, remnants of hemoglobin genes can be found in their genome.[9]
Vertebrate red blood cells consist mainly of hemoglobin, a complex metalloprotein containing heme groups whose iron atoms temporarily bind to oxygen molecules (O2) in the lungs or gills and release them throughout the body. Oxygen can easily diffuse through the red blood cell's cell membrane. Hemoglobin in the red blood cells also carries some of the waste product carbon dioxide back from the tissues; most waste carbon dioxide, however, is transported back to the pulmonary capillaries of the lungs as bicarbonate (HCO3−) dissolved in the blood plasma. Myoglobin, a compound related to hemoglobin, acts to store oxygen in muscle cells.[10]
The color of red blood cells is due to the heme group of hemoglobin. The blood plasma alone is straw-colored, but the red blood cells change color depending on the state of the hemoglobin: when combined with oxygen the resulting oxyhemoglobin is scarlet, and when oxygen has been released the resulting deoxyhemoglobin is of a dark red burgundy color. However, blood can appear bluish when seen through the vessel wall and skin.[11] Pulse oximetry takes advantage of the hemoglobin color change to directly measure the arterial blood oxygen saturation using colorimetric techniques. Hemoglobin also has a very high affinity for carbon monoxide, forming carboxyhemoglobin which is a very bright red in color. Flushed, confused patients with a saturation reading of 100% on pulse oximetry are sometimes found to be suffering from carbon monoxide poisoning.
Having oxygen-carrying proteins inside specialized cells (as opposed to oxygen carriers being dissolved in body fluid) was an important step in the evolution of vertebrates as it allows for less viscous blood, higher concentrations of oxygen, and better diffusion of oxygen from the blood to the tissues. The size of red blood cells varies widely among vertebrate species; red blood cell width is on average about 25% larger than capillary diameter, and it has been hypothesized that this improves the oxygen transfer from red blood cells to tissues.[12]
Mammals
Typical mammalian red blood cells: (a) seen from surface; (b) in profile, forming rouleaux; (c) rendered spherical by water; (d) rendered crenate by salt. (c) and (d) do not normally occur in the body.
The red blood cells of mammals are typically shaped as biconcave disks: flattened and depressed in the center, with a dumbbell-shaped cross section, and a torus-shaped rim on the edge of the disk. This distinctive biconcave shape optimises the flow properties of blood in the large vessels, such as maximization of laminar flow and minimization of platelet scatter, which suppresses their atherogenic activity in those large vessels.[13] Moreover, this shape allows for a high surface-area-to-volume (SA/V) ratio to facilitate diffusion of gases.[14] However, there are some exceptions concerning shape in the artiodactyl order (even-toed ungulates including cattle, deer, and their relatives), which displays a wide variety of bizarre red blood cell morphologies: small and highly ovaloid cells in llamas and camels (family Camelidae), tiny spherical cells in mouse deer (family Tragulidae), and cells which assume fusiform, lanceolate, crescentic, and irregularly polygonal and other angular forms in red deer and wapiti (family Cervidae). Members of this order have clearly evolved a mode of red blood cell development substantially different from the mammalian norm.[7][15] Overall, mammalian red blood cells are remarkably flexible and deformable so as to squeeze through tiny capillaries, as well as to maximize their apposing surface by assuming a cigar shape, where they efficiently release their oxygen load.[16]
Red blood cells in mammals are unique amongst vertebrates as they do not have nuclei when mature. Red blood cells of mammals cells have nuclei during early phases of erythropoiesis, but extrude them during development as they mature in order to provide more space for hemoglobin. The red blood cells without nuclei, called reticulocytes, go on to lose all other cellular organelles such as their mitochondria, Golgi apparatus and endoplasmic reticulum.
The spleen acts as a reservoir of red blood cells, but this effect is somewhat limited in humans. In some other mammals such as dogs and horses, the spleen sequesters large numbers of red blood cells which are dumped into the blood during times of exertion stress, yielding a higher oxygen transport capacity.
Scanning electron micrograph of blood cells. From left to right: human red blood cell, thrombocyte (platelet), leukocyte.
Human
Two drops of blood are shown with a bright red oxygenated drop on the left and a deoxygenated drop on the right.
Animation of a typical human red blood cell cycle in the circulatory system. This animation occurs at a faster rate (~20 seconds of the average 60-second cycle) and shows the red blood cell deforming as it enters capillaries, as well as the bars changing color as the cell alternates in states of oxygenation along the circulatory system.
A typical human red blood cell has a disk diameter of approximately 6.2–8.2 µm[17] and a thickness at the thickest point of 2–2.5 µm and a minimum thickness in the centre of 0.8–1 µm, being much smaller than most other human cells. These cells have an average volume of about 90 fL[18] with a surface of about 136 μm2, and can swell up to a sphere shape containing 150 fL, without membrane distension.
Adult humans have roughly 20–30 trillion red blood cells at any given time, constituting approximately 70% all cells by number.[19] Women have about 4–5 million red blood cells per microliter (cubic millimeter) of blood and men about 5–6 million; people living at high altitudes with low oxygen tension will have more. Red blood cells are thus much more common than the other blood particles: there are about 4,000–11,000 white blood cells and about 150,000–400,000 platelets per microliter.
Human red blood cells take on average 60 seconds to complete one cycle of circulation.[3][5][20]
The blood's red color is due to the spectral properties of the hemic iron ions in hemoglobin. Each human red blood cell contains approximately 270 million[citation needed] of these hemoglobin molecules. Each hemoglobin molecule carries four heme groups; hemoglobin constitutes about a third of the total cell volume. Hemoglobin is responsible for the transport of more than 98% of the oxygen in the body (the remaining oxygen is carried dissolved in the blood plasma). The red blood cells of an average adult human male store collectively about 2.5 grams of iron, representing about 65% of the total iron contained in the body.[21][22]
Microstructure
Nucleus
Red blood cells in mammals anucleate when mature, meaning that they lack a cell nucleus. In comparison, the red blood cells of other vertebrates have nuclei; the only known exceptions are salamanders of the Batrachoseps genus and fish of the Maurolicus genus with closely related species.[23][24]
The elimination of the nucleus in vertebrate red blood cells has been offered as an explanation for the subsequent accumulation of non-coding DNA in the genome.[25] The argument runs as follows: Efficient gas transport requires red blood cells to pass through very narrow capillaries, and this constrains their size. In the absence of nuclear elimination, the accumulation of repeat sequences is constrained by the volume occupied by the nucleus, which increases with genome size.
Nucleated red blood cells in mammals consist of two forms: normoblasts, which are normal erythropoietic precursors to mature red blood cells, and megaloblasts, which are abnormally large precursors that occur in megaloblastic anemias.
Membrane composition
Red blood cells are deformable, flexible, are able to adhere to other cells, and are able to interface with immune cells. Their membrane plays many roles in this. These functions are highly dependent on the membrane composition. The red blood cell membrane is composed of 3 layers: the glycocalyx on the exterior, which is rich in carbohydrates; the lipid bilayer which contains many transmembrane proteins, besides its lipidic main constituents; and the membrane skeleton, a structural network of proteins located on the inner surface of the lipid bilayer. Half of the membrane mass in human and most mammalian red blood cells are proteins. The other half are lipids, namely phospholipids and cholesterol.[26]
Membrane lipids
The most common red blood cell cell membrane lipids, schematically disposed as they are distributed on the bilayer. Relative abundances are not at scale.
The red blood cell cell membrane comprises a typical lipid bilayer, similar to what can be found in virtually all human cells. Simply put, this lipid bilayer is composed of cholesterol and phospholipids in equal proportions by weight. The lipid composition is important as it defines many physical properties such as membrane permeability and fluidity. Additionally, the activity of many membrane proteins is regulated by interactions with lipids in the bilayer.
Unlike cholesterol, which is evenly distributed between the inner and outer leaflets, the 5 major phospholipids are asymmetrically disposed, as shown below:
Outer monolayer
Phosphatidylcholine (PC);
Sphingomyelin (SM).
Inner monolayer
Phosphatidylethanolamine (PE);
Phosphoinositol (PI) (small amounts).
Phosphatidylserine (PS);
This asymmetric phospholipid distribution among the bilayer is the result of the function of several energy-dependent and energy-independent phospholipid transport proteins. Proteins called “Flippases” move phospholipids from the outer to the inner monolayer, while others called “floppases” do the opposite operation, against a concentration gradient in an energy dependent manner. Additionally, there are also “scramblase” proteins that move phospholipids in both directions at the same time, down their concentration gradients in an energy independent manner. There is still considerable debate ongoing regarding the identity of these membrane maintenance proteins in the red cell membrane.
The maintenance of an asymmetric phospholipid distribution in the bilayer (such as an exclusive localization of PS and PIs in the inner monolayer) is critical for the cell integrity and function due to several reasons:
Macrophages recognize and phagocytose red cells that expose PS at their outer surface. Thus the confinement of PS in the inner monolayer is essential if the cell is to survive its frequent encounters with macrophages of the reticuloendothelial system, especially in the spleen.
Premature destruction of thallassemic and sickle red cells has been linked to disruptions of lipid asymmetry leading to exposure of PS on the outer monolayer.
An exposure of PS can potentiate adhesion of red cells to vascular endothelial cells, effectively preventing normal transit through the microvasculature. Thus it is important that PS is maintained only in the inner leaflet of the bilayer to ensure normal blood flow in microcirculation.
Both PS and phosphatidylinositol-4,5-bisphosphate (PIP2) can regulate membrane mechanical function, due to their interactions with skeletal proteins such as spectrin and protein 4.1R. Recent studies have shown that binding of spectrin to PS promotes membrane mechanical stability. PIP2 enhances the binding of protein band 4.1R to glycophorin C but decreases its interaction with protein band 3, and thereby may modulate the linkage of the bilayer to the membrane skeleton.
The presence of specialized structures named "lipid rafts" in the red blood cell membrane have been described by recent studies. These are structures enriched in cholesterol and sphingolipids associated with specific membrane proteins, namely flotillins, stomatins (band 7), G-proteins, and β-adrenergic receptors. Lipid rafts that have been implicated in cell signaling events in nonerythroid cells have been shown in erythroid cells to mediate β2-adregenic receptor signaling and increase cAMP levels, and thus regulating entry of malarial parasites into normal red cells.[27][28]
Membrane proteins
Red blood cell membrane proteins separated by SDS-PAGE and silverstained[29]
The proteins of the membrane skeleton are responsible for the deformability, flexibility and durability of the red blood cell, enabling it to squeeze through capillaries less than half the diameter of the red blood cell (7–8 μm) and recovering the discoid shape as soon as these cells stop receiving compressive forces, in a similar fashion to an object made of rubber.
There are currently more than 50 known membrane proteins, which can exist in a few hundred up to a million copies per red blood cell. Approximately 25 of these membrane proteins carry the various blood group antigens, such as the A, B and Rh antigens, among many others. These membrane proteins can perform a wide diversity of functions, such as transporting ions and molecules across the red cell membrane, adhesion and interaction with other cells such as endothelial cells, as signaling receptors, as well as other currently unknown functions. The blood types of humans are due to variations in surface glycoproteins of red blood cells. Disorders of the proteins in these membranes are associated with many disorders, such as hereditary spherocytosis, hereditary elliptocytosis, hereditary stomatocytosis, and paroxysmal nocturnal hemoglobinuria.[26][27]
The red blood cell membrane proteins organized according to their function:
Red blood cell membrane major proteins
Transport
Band 3 – Anion transporter, also an important structural component of the red blood cell cell membrane, makes up to 25% of the cell membrane surface, each red cell contains approximately one million copies. Defines the Diego Blood Group;[30]
Aquaporin 1 – water transporter, defines the Colton Blood Group;
Glut1 – glucose and L-dehydroascorbic acid transporter;
Kidd antigen protein – urea transporter;
RhAG – gas transporter, probably of carbon dioxide, defines Rh Blood Group and the associated unusual blood group phenotype Rhnull;
Na+/K+ – ATPase;
Ca2+ – ATPase;
Na+ K+ 2Cl− – cotransporter;
Na+-Cl− – cotransporter;
Na-H exchanger;
K-Cl – cotransporter;
Gardos Channel.
Cell adhesion
ICAM-4 – interacts with integrins;
BCAM – a glycoprotein that defines the Lutheran blood group and also known as Lu or laminin-binding protein.
Structural role – The following membrane proteins establish linkages with skeletal proteins and may play an important role in regulating cohesion between the lipid bilayer and membrane skeleton, likely enabling the red cell to maintain its favorable membrane surface area by preventing the membrane from collapsing (vesiculating).
Ankyrin-based macromolecular complex – proteins linking the bilayer to the membrane skeleton through the interaction of their cytoplasmic domains with Ankyrin.
Band 3 – also assembles various glycolytic enzymes, the presumptive CO2 transporter, and carbonic anhydrase into a macromolecular complex termed a "metabolon," which may play a key role in regulating red cell metabolism and ion and gas transport function);
RhAG – also involved in transport, defines associated unusual blood group phenotype Rhmod.
Protein 4.1R-based macromolecular complex – proteins interacting with Protein 4.1R.
Protein 4.1R – weak expression of Gerbich antigens;
Glycophorin C and D – glycoprotein, defines Gerbich Blood Group;
XK – defines the Kell Blood Group and the Mcleod unusual phenotype (lack of Kx antigen and greatly reduced expression of Kell antigens);
RhD/RhCE – defines Rh Blood Group and the associated unusual blood group phenotype Rhnull;
Duffy protein – has been proposed to be associated with chemokine clearance;[31]
Adducin – interaction with band 3;
Dematin- interaction with the Glut1 glucose transporter.
[26][27]
Surface electrostatic potential
The zeta potential is an electrochemical property of cell surfaces that is determined by the net electrical charge of molecules exposed at the surface of cell membranes of the cell. The normal zeta potential of the red blood cell is −15.7 millivolts (mV).[32] Much of this potential appears to be contributed by the exposed sialic acid residues in the membrane: their removal results in zeta potential of −6.06 mV.
Function
Secondary functions
When red blood cells undergo shear stress in constricted vessels, they release ATP, which causes the vessel walls to relax and dilate so as to promote normal blood flow.[33]
When their hemoglobin molecules are deoxygenated, red blood cells release S-nitrosothiols, which also act to dilate blood vessels,[34] thus directing more blood to areas of the body depleted of oxygen.
Red blood cells can also synthesize nitric oxide enzymatically, using L-arginine as substrate, as do endothelial cells.[35] Exposure of red blood cells to physiological levels of shear stress activates nitric oxide synthase and export of nitric oxide,[36] which may contribute to the regulation of vascular tonus.
Red blood cells can also produce hydrogen sulfide, a signalling gas that acts to relax vessel walls. It is believed that the cardioprotective effects of garlic are due to red blood cells converting its sulfur compounds into hydrogen sulfide.[37]
Red blood cells also play a part in the body's immune response: when lysed by pathogens such as bacteria, their hemoglobin releases free radicals, which break down the pathogen's cell wall and membrane, killing it.[38][39]
Cellular processes
As a result of not containing mitochondria, red blood cells use none of the oxygen they transport; instead they produce the energy carrier ATP by the glycolysis of glucose and lactic acid fermentation on the resulting pyruvate.[40][41] Furthermore the pentose phosphate pathway plays an important role in red blood cells; see glucose-6-phosphate dehydrogenase deficiency for more.
As red blood cells contain no nucleus, protein biosynthesis is currently assumed to be absent in these cells.
Because of the lack of nuclei and organelles, mature red blood cells do not contain DNA and cannot synthesize any RNA, and consequently cannot divide and have limited repair capabilities.[42] The inability to carry out protein synthesis means that no virus can evolve to target mammalian red blood cells.[43] However, infection with parvoviruses (such as human parvovirus B19) can affect erythroid precursors, as recognized by the presence of giant pronormoblasts with viral particles and inclusion bodies, thus temporarily depleting the blood of reticulocytes and causing anemia.[44]
Life cycle
Human red blood cells are produced through a process named erythropoiesis, developing from committed stem cells to mature red blood cells in about 7 days. When matured, in a healthy individual these cells live in blood circulation for about 100 to 120 days (and 80 to 90 days in a full term infant).[45] At the end of their lifespan, they are removed from circulation. In many chronic diseases, the lifespan of the red blood cells is reduced.
Creation
Erythropoiesis is the process by which new red blood cells are produced; it lasts about 7 days. Through this process red blood cells are continuously produced in the red bone marrow of large bones. (In the embryo, the liver is the main site of red blood cell production.) The production can be stimulated by the hormone erythropoietin (EPO), synthesised by the kidney. Just before and after leaving the bone marrow, the developing cells are known as reticulocytes; these constitute about 1% of circulating red blood cells.
Functional lifetime
The functional lifetime of an red blood cell is about 100–120 days, during which time the red blood cells are continually moved by the blood flow push (in arteries), pull (in veins) and a combination of the two as they squeeze through microvessels such as capillaries. They are also recycled in the bone marrow.[46]
Senescence
The aging red blood cell undergoes changes in its plasma membrane, making it susceptible to selective recognition by macrophages and subsequent phagocytosis in the mononuclear phagocyte system (spleen, liver and lymph nodes), thus removing old and defective cells and continually purging the blood. This process is termed eryptosis, red blood cell programmed cell death.[47] This process normally occurs at the same rate of production by erythropoiesis, balancing the total circulating red blood cell count. Eryptosis is increased in a wide variety of diseases including sepsis, haemolytic uremic syndrome, malaria, sickle cell anemia, beta-thalassemia, glucose-6-phosphate dehydrogenase deficiency, phosphate depletion, iron deficiency and Wilson's disease. Eryptosis can be elicited by osmotic shock, oxidative stress, energy depletion as well as a wide variety of endogenous mediators and xenobiotics. Excessive eryptosis is observed in red blood cells lacking the cGMP-dependent protein kinase type I or the AMP-activated protein kinase AMPK. Inhibitors of eryptosis include erythropoietin, nitric oxide, catecholamines and high concentrations of urea.
Much of the resulting breakdown products are recirculated in the body. The heme constituent of hemoglobin are broken down into iron (Fe3+) and biliverdin. The biliverdin is reduced to bilirubin, which is released into the plasma and recirculated to the liver bound to albumin. The iron is released into the plasma to be recirculated by a carrier protein called transferrin. Almost all red blood cells are removed in this manner from the circulation before they are old enough to hemolyze. Hemolyzed hemoglobin is bound to a protein in plasma called haptoglobin, which is not excreted by the kidney.[48]
Clinical significance
Disease
Affected by Sickle-cell disease, red blood cells alter shape and threaten to damage internal organs.
Blood diseases involving the red blood cells include:
Anemias (or anaemias) are diseases characterized by low oxygen transport capacity of the blood, because of low red cell count or some abnormality of the red blood cells or the hemoglobin.
Iron deficiency anemia is the most common anemia; it occurs when the dietary intake or absorption of iron is insufficient, and hemoglobin, which contains iron, cannot be formed
Sickle-cell disease is a genetic disease that results in abnormal hemoglobin molecules. When these release their oxygen load in the tissues, they become insoluble, leading to mis-shaped red blood cells. These sickle shaped red cells are less deformable and viscoelastic meaning that they have become rigid and can cause blood vessel blockage, pain, strokes, and other tissue damage.
Thalassemia is a genetic disease that results in the production of an abnormal ratio of hemoglobin subunits.
Hereditary spherocytosis syndromes are a group of inherited disorders characterized by defects in the red blood cell's cell membrane, causing the cells to be small, sphere-shaped, and fragile instead of donut-shaped and flexible. These abnormal red blood cells are destroyed by the spleen. Several other hereditary disorders of the red blood cell membrane are known.[49]
Pernicious anemia is an autoimmune disease wherein the body lacks intrinsic factor, required to absorb vitamin B12 from food. Vitamin B12 is needed for the production of hemoglobin.
Aplastic anemia is caused by the inability of the bone marrow to produce blood cells.
Pure red cell aplasia is caused by the inability of the bone marrow to produce only red blood cells.
Effect of osmotic pressure on blood cells
Micrographs of the effects of osmotic pressure
Hemolysis is the general term for excessive breakdown of red blood cells. It can have several causes and can result in hemolytic anemia.
The malaria parasite spends part of its life-cycle in red blood cells, feeds on their hemoglobin and then breaks them apart, causing fever. Both sickle-cell disease and thalassemia are more common in malaria areas, because these mutations convey some protection against the parasite.
Polycythemias (or erythrocytoses) are diseases characterized by a surplus of red blood cells. The increased viscosity of the blood can cause a number of symptoms.
In polycythemia vera the increased number of red blood cells results from an abnormality in the bone marrow.
Several microangiopathic diseases, including disseminated intravascular coagulation and thrombotic microangiopathies, present with pathognomonic (diagnostic) red blood cell fragments called schistocytes. These pathologies generate fibrin strands that sever red blood cells as they try to move past a thrombus.
Transfusion
Main article: Blood transfusion
Red blood cells may be given as part of a blood transfusion. Blood may be donated from another person, or stored by the recipient at an earlier date. Donated blood usually requires screening to ensure that donors do not contain risk factors for the presence of blood-borne diseases, or will not suffer themselves by giving blood. Blood is usually collected and tested for common or serious Blood-borne diseases including Hepatitis B, Hepatitis C and HIV. The blood type (A, B, AB, or O) or the blood product is identified. This relates to the presence of antigens on the cell's surface. After this process, the blood is stored, and within a short duration is used. Blood can be given as a whole product or the red blood cells separated as packed red blood cells.
Blood is often transfused when there is known anaemia, active bleeding, or when there is an expectation of serious blood loss, such as prior to an operation. Before blood is given, a small sample of the recipient's blood is tested with the transfusion in a process known as cross-matching. In addition to the transmission of infection, certain types of transfusion reaction.
In 2008 it was reported that human embryonic stem cells had been successfully coaxed into becoming red blood cells in the lab. The difficult step was to induce the cells to eject their nucleus; this was achieved by growing the cells on stromal cells from the bone marrow. It is hoped that these artificial red blood cells can eventually be used for blood transfusions.[50]
Tests
Several blood tests involve red blood cells. These include a RBC count (the number of red blood cells per volume of blood), calculation of the hematocrit (percentage of blood volume occupied by red blood cells), and the erythrocyte sedimentation rate. The blood type needs to be determined to prepare for a blood transfusion or an organ transplantation.
Many diseases involving red blood cells are diagnosed with a blood film (or peripheral blood smear), where a thin layer of blood is smeared on a microscope slide. This may reveal abnormalities of red blood cell shape and form. When red blood cells sometimes occur as a stack, flat side next to flat side. This is known as rouleaux formation, and it occurs more often if the levels of certain serum proteins are elevated, as for instance during inflammation.
Separation and blood doping
Red blood cells can be obtained from whole blood by centrifugation, which separates the cells from the blood plasma in a process known as blood fractionation. Packed red blood cells, which are made in this way from whole blood with the plasma removed, are used in transfusion medicine.[51] During plasma donation, the red blood cells are pumped back into the body right away and only the plasma is collected.
Some athletes have tried to improve their performance by blood doping: first about 1 litre of their blood is extracted, then the red blood cells are isolated, frozen and stored, to be reinjected shortly before the competition. (Red blood cells can be conserved for 5 weeks at −79 °C or −110 °F) This practice is hard to detect but may endanger the human cardiovascular system which is not equipped to deal with blood of the resulting higher viscosity. Another method of blood doping involves injection with erythropoietin in order to stimulate production of red blood cells. Both practices are banned by the World Anti-Doping Agency.
History
The first person to describe red blood cells was the young Dutch biologist Jan Swammerdam, who had used an early microscope in 1658 to study the blood of a frog.[52] Unaware of this work, Anton van Leeuwenhoek provided another microscopic description in 1674, this time providing a more precise description of red blood cells, even approximating their size, "25,000 times smaller than a fine grain of sand".
In 1901, Karl Landsteiner published his discovery of the three main blood groups—A, B, and C (which he later renamed to O). Landsteiner described the regular patterns in which reactions occurred when serum was mixed with red blood cells, thus identifying compatible and conflicting combinations between these blood groups. A year later Alfred von Decastello and Adriano Sturli, two colleagues of Landsteiner, identified a fourth blood group—AB.
In 1959, by use of X-ray crystallography, Dr. Max Perutz was able to unravel the structure of hemoglobin, the red blood cell protein that carries oxygen.[53]
The oldest intact red blood cells ever discovered were found in Ötzi the Iceman, a natural mummy of a man who died around 3255 BCE. These cells were discovered in May 2012.[54]
See also
Altitude training
Blood serum
Erythrocyte deformability
Erythrocyte fragility
Blood substitute
Packed red blood cells
Red blood cell indices
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External links
Wikimedia Commons has media related to Red blood cells.
Blood Groups and Red Cell Antigens by Laura Dean. Searchable and downloadable online textbook in the public domain.
Database of vertebrate erythrocyte sizes.
Red Gold, PBS site containing facts and history
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GO Transit fares
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GO Transit is the inter-regional transportation authority of the Greater Toronto and Hamilton Area. It is Canada's oldest regional transit system, first serving passengers in 1967.
Contents
[hide]
1 Fare zones
1.1 List of fare zones
2 Fare classes
3 Fare types
3.1 Presto card
3.2 Fare integration
3.2.1 Via Rail integration
3.2.2 Local transit integration
3.2.3 GO Transit co-fares
3.2.4 Other local transit integration
4 References
5 External links
Fare zones[edit]
The service area of GO Transit is divided into a number of fare zones, each of which belonging to a number of fare corridors. Fares on GO Transit are based on the distance between originating and destination fare zones. Tickets between two fare zones are valid for travel between the two fare zones stated on the ticket, as well as between any two fare zones that lie between them on the same fare corridor. Tickets are not generally issued between two fare zones on different corridors, with the following notable exceptions:
Due to all-day GO train service and the interlining of the Lakeshore West line and the Lakeshore East line, tickets may be issued for trips starting in a fare zone on one line and ending in a fare zone on the other. Similarly, due to the number of GO bus connections between stations on the Lakeshore West line and the Milton line (including Square One Bus Terminal, but excluding Lisgar GO station), tickets may be issued for trips starting in a fare zone in Mississauga or Milton outside of the Lakeshore West line and fare zones on either of the Lakeshore lines.
Tickets are generally issued for trips between a fare zone inside the City of Toronto and a fare zone outside the City of Toronto, even if they lie in different fare corridors; such tickets are valid between the fare zone outside the City of Toronto to any fare zone inside the City of Toronto for which the ticket value is equal or less. (For example, a ticket from Square One Bus Terminal to Scarborough Centre Bus Terminal would also be valid for trips to Kennedy GO Station, as the trips are of equal value, even though all three locations are on different fare corridors.) Note that Milliken GO Station, despite physically located within the City of Toronto, is considered to be in a fare zone outside of the City of Toronto for historical reasons.[citation needed]
As all GO Transit tickets are only valid for continuous trips of no more than two and a half hours, multiple tickets may be needed for longer trips.
List of fare zones[edit]
Fare zones 1–9, 19, 77, 89, and 90 (shaded in green) are located within the City of Toronto.
No.
Name
Stops
Corridor
1 Toronto Coach Terminal No longer used by GO Transit
2 Toronto Union Station
Union Station Bus Terminal
Exhibition GO Station
Bloor GO Station (all corridors except Highway 407)
3 Etobicoke South Mimico GO Station
Long Branch GO Station
Kipling GO Station Lakeshore West line
Milton line
Niagara Falls
4 Metro Northwest Weston GO Station
Etobicoke North GO Station
Humber College Milton line
Kitchener line
Bolton
5 Metro North Oriole GO Station
Old Cummer GO Station
Yorkdale Bus Terminal
York Mills Bus Terminal
Finch Bus Terminal
Sheppard-Yonge subway station Kitchener line
Beaverton (via Newmarket)
Richmond Hill line
Barrie line
Lakeshore East line
Bolton
6 Scarborough South Danforth GO Station
Scarborough GO Station
Eglinton GO Station Beaverton (via Whitby)
Lakeshore East line
Peterborough
7 Scarborough Centre Agincourt GO Station
Scarborough Centre Bus Terminal Beaverton (via Whitby)
Stouffville line
Lakeshore East line
Peterborough
8 Guildwood Guildwood GO Station Beaverton (via Whitby)
Lakeshore East line
Peterborough
9 Rouge Hill Rouge Hill GO Station Beaverton (via Whitby)
Lakeshore East line
Peterborough
11 Port Credit—Cooksville Port Credit GO Station
Dixie GO Station
Cooksville GO Station Lakeshore West line
Milton line
Niagara Falls
12 Clarkson—Erindale Clarkson GO Station
Erindale GO Station Lakeshore West line
Milton line
Niagara Falls
13 Oakville Oakville GO Station
Sheridan College Lakeshore West line
Highway 407
Niagara Falls
14 Bronte Bronte GO Station
Bronte Carpool Lot Lakeshore West line
Highway 407
Niagara Falls
15 Appleby Appleby GO Station Lakeshore West line
Niagara Falls
16 Burlington Burlington GO Station
Burlington Carpool Lot Lakeshore West line
Highway 407
Niagara Falls
17 Aldershot Aldershot GO Station Lakeshore West line
18 Hamilton Hamilton GO Centre
McMaster University Bus Terminal
West Harbour GO Station Lakeshore West line
Highway 407
19 York University York University GO Station
York University Bus Loop Beaverton (via Newmarket)
Barrie line
Highway 407
20 Square One Square One Bus Terminal Milton line
Highway 407
21 Streetsville Streetsville GO Station Milton line
22 Meadowvale Meadowvale GO Station
Meadowvale Town Centre
Meadowvale Business Park
Hurontario at 407 GO Park & Ride Milton line
Highway 407
23 Lisgar Lisgar GO Station Milton line
24 Milton Milton GO Station
Milton GO Park & Ride Milton line
26 Cambridge SmartCentres iXpress Station Milton line
27 Kitchener Kitchener railway station
Charles St. Transit Terminal
Wilfrid Laurier University (Laurier iXpress Station)
University of Waterloo (UW Davis iXpress Station) Kitchener line
Milton line
28 Pontypool Clarington North GO Park & Ride Peterborough
29 Millbrook Cavan/Millbrook GO Park & Ride Peterborough
30 Peterborough Peterborough Transit Terminal
Trent University
Peterborough GO Park & Ride Peterborough
31 Malton Malton GO Station
Toronto Pearson International Airport Kitchener line
Orangeville
32 Bramalea Bramalea GO Station
Bramalea City Centre Terminal
Trinity Common Bus Terminal Kitchener line
Orangeville
Highway 407
33 Brampton Brampton GO Station
Brampton Bus Terminal
Shoppers World Terminal Kitchener line
Orangeville
34 Mount Pleasant Mount Pleasant GO Station Kitchener line
35 Georgetown Georgetown GO Station Kitchener line
36 Silvercreek Kitchener line
37 Acton Acton GO Station Kitchener line
38 Rockwood Kitchener line
39 Guelph Guelph Central GO Station
Guelph Bus Terminal
University of Guelph
Aberfoyle GO Park and Ride Kitchener line
Highway 407
41 Victoria Orangeville
42 Caledon Orangeville
43 Orangeville Orangeville Mall
Orangeville GO Park & Ride Orangeville
44 East Gwillimbury East Gwillimbury GO Station Beaverton (via Newmarket)
Barrie line
45 Queensville Beaverton (via Newmarket)
46 Keswick Beaverton (via Newmarket)
47 Island Grove Beaverton (via Newmarket)
48 Sutton Beaverton (via Newmarket)
49 Virginia Beaverton (via Newmarket)
51 Brooklin—Myrtle Beaverton (via Whitby)
Highway 407
52 Port Perry Beaverton (via Whitby)
53 Saintfield Beaverton (via Whitby)
54 Sunderland Beaverton (via Whitby)
55 Beaverton Beaverton (via Newmarket)
Beaverton (via Whitby)
56 Kleinburg Bolton
57 Nobleton Bolton
58 Bolton Bolton
60 Langstaff Langstaff GO Station
Richmond Hill Centre Terminal Beaverton (via Newmarket)
Richmond Hill line
Barrie line
Highway 407
61 Richmond Hill—Maple—Rutherford Rutherford GO Station
Maple GO Station
Richmond Hill GO Station
Wonderland Terminal
Major Mackenzie at Highway 400 Carpool Lot Beaverton (via Newmarket)
Richmond Hill line
Barrie line
62 Oak Ridges—King King City GO Station Beaverton (via Newmarket)
Barrie line
63 Aurora Aurora GO Station
Aurora Carpool Lot Beaverton (via Newmarket)
Barrie line
64 Newmarket Newmarket GO Station
Newmarket Bus Terminal
Highway 9 at 400 Carpool Lot Barrie line
65 Bradford Bradford GO Station Barrie line
66 Deerhurst Barrie line
67 Churchill Barrie line
68 Barrie South—Stroud Barrie South GO Station Barrie line
69 Barrie Allandale Waterfront GO Station
Barrie Bus Terminal Barrie line
70 Milliken Milliken GO Station Stouffville line
71 Unionville Unionville GO Station Stouffville line
Highway 407
72 Markham—Centennial Centennial GO Station
Markham GO Station Stouffville line
Highway 407
73 Mount Joy—Wideman Mount Joy GO Station Stouffville line
Highway 407
74 Stouffville—Lincolnville Stouffville GO Station
Lincolnville GO Station Stouffville line
75 Goodwood Stouffville line
76 Uxbridge Stouffville line
77 Kennedy Kennedy GO Station Stouffville line
80 Stoney Creek Stoney Creek GO Park & Ride Niagara Falls
81 Grimsby Grimsby GO Park & Ride Niagara Falls
82 Lincoln Niagara Falls
83 St. Catharines St. Catharines railway station
Fairview Mall Niagara Falls
84 Niagara Falls Niagara Falls railway station
Niagara Falls Transit Terminal Niagara Falls
86 Brougham Highway 407
87 Greenwood Highway 407
88 Durham College Durham College
University of Ontario Institute of Technology Lakeshore East line
Highway 407
89 Malvern Centennial College
University of Toronto Scarborough Highway 407
90 North Rouge Lakeshore East line
91 Pickering Pickering GO Station Beaverton (via Whitby)
Lakeshore East line
Highway 407
Peterborough
92 Ajax Ajax GO Station
Ajax Bus Terminal Beaverton (via Whitby)
Lakeshore East line
Peterborough
93 Whitby Whitby GO Station Beaverton (via Whitby)
Lakeshore East line
Peterborough
94 Oshawa Oshawa GO Station
Oshawa Bus Terminal Lakeshore East line
Peterborough
95 Courtice Lakeshore East line
96 Bowmanville Bowmanville GO Park & Ride Lakeshore East line
97 Newcastle Newcastle Carpool Lot Lakeshore East line
Peterborough
Fare classes[edit]
GO Transit offers four passenger fare classes: adult, senior, child, and student. Senior fares are available for passengers over the age of 65, while child fares are available for passengers between the ages of 6 and 12. Passengers between the ages of 1 and 5 ride free with an accompanying adult, but each adult may only ride with one child for free in this manner; any unaccompanied child must pay child fares. Passengers under the age of 1 always ride free on GO Transit.
Student fares are only available for those with acceptable student identification:
For elementary and secondary school students, school-issued student identification is needed. Note that these do not necessarily need to be identification issued by a school within GO Transit's service area.
Full-time students of post-secondary institutions anywhere in Canada (not necessarily within GO Transit's service area) for which student identification is reissued each year may use their school-issued student identification for proof of student fare qualification.
Students at Durham College, University of Ontario Institute of Technology, and Trent University's Oshawa Campus require their school-issued student identification for proof of student fare qualification.
Students at Centennial College, Humber College, Michener Institute, Mohawk College, Niagara University, Sheridan College, and Trent University's Peterborough campus require a sticker to be affixed to their school-issued student identification for proof of student fare qualification.
Students at McMaster University holding a Hamilton Street Railway U-Pass require their school-issued student identification for proof of student fare qualification.
All other post-secondary students require the GO Transit student ID. It is valid for full-time students from October of one year to October of the next year, as long as the student retains full-time status. GO Transit student ID application forms may be obtained from the registrar's office of their institutions. These must be processed at GO Transit's offices at Union Station before they may be used as proof of student fare qualification. Students at Seneca College, Ryerson University, York University, University of Waterloo, and the University of Guelph may also obtain GO Transit student identification online, while students at the University of Toronto's St. George, Mississauga, and Scarborough campuses may obtain their GO Transit student identification on campus.
Fare types[edit]
Up until July 31, 2012, GO Transit offered five types of fares:
Single-Ride Tickets are for one single journey between two fare zones. These expire four hours after purchase, and are not refundable.
Two-Ride Tickets are for two single journeys between two fare zones. These do not expire, but are not refundable.
Day Passes are for unlimited journeys between two fare zones from 3AM one day to 3AM the next. These are not refundable.
Ten-Ride Tickets are for 10 single journeys between two fare zones. These do not expire, and unused portions may be refunded 30 days after purchase.
Monthly Passes are for unlimited journeys between two fare zones for one calendar month. They are available for purchase 10 days before the month to 14 days into the month, and must be signed by its user for it to be valid. These may be refundable.
GO Transit stopped selling two-ride and ten-ride tickets on June 1, 2012 and stopped accepting two-ride and ten-ride tickets on July 31, 2012, in favour of using Presto Cards exclusively; paper monthly passes will be discontinued effective 1 January 2013;[1] day passes remain available for purchase. Passengers may convert any unused rides on their tickets to equivalent value on a Presto Card before the end of 2012.
There is no distinction between bus and train service, with the exception of the seasonal train service to Niagara Falls: passengers paying single-ride tickets and travelling by train to Niagara Falls are assessed a $3 surcharge on top of existing fares.
Fares between two fare zones for a particular type of ticket and passenger category are based on multiples of the standard (adult single-ride) fare as follows:
Fare multiples for ticket users
Adult
Senior/Child
Student
Single-Ride 1 0.5 -
Day Pass 2 1 -
Monthly Pass 34 20 28
Fares are rounded to the nearest $0.05 for single-ride fares and day passes and $1.00 for monthly passes, after fare multiples are applied. The minimum adult single-ride fare (for travel on GO Transit within one fare zone) is always $4.50; other fares depend on the fare zones travelled.
Presto card[edit]
PRESTO card reader at Oshawa station
The Presto card is also accepted on GO Transit. Passengers using the Presto Card must tap their cards against the reader upon embarking and disembarking the train or bus. Upon embarking, the minimum fare ($5.30) is always deducted from the card, with fare adjustments performed as the passenger disembarks.
Failure to tap the card upon disembarking ("missed tap-off") will cause the maximum fare to be deducted, under the assumption that the passenger travelled to the point furthest away from the origin point. (For example, passengers failing to tap off when boarding at Union Station will be assessed the fare travelling from Union Station to Kitchener GO Station, the point furthest away on any GO train route serving Union Station.) Passengers regularly travel by GO train may also set a "default trip" between two GO train stations, allowing them to avoid tapping their cards upon disembarking; passengers doing so will have the appropriate fare deducted upon embarking at either endpoint.
Adult and student passengers using the Presto card receive a further fare discount, depending on the number of trips that have been made per month between two fare zones.
Fare multiples for Presto Card users
Adult
Student
Child/Senior
First 30 trips 0.8885 0.816 0.4735
Trips 31–35 0.8885 0.05 0.4735
Trips 36–40 0.1225 0.05 0.4735
Each trip thereafter Free Free Free
If there are rides between different sets of fare zones, the discounts after the 30th trip (students) and 35th trip (adults) will be based on the total value of rides taken so far; the fare discounts are structured so that adult and student passengers pay less over the course of a month compared to paying for a paper monthly pass ahead of time; the discrepancy is to encourage passengers to adopt the Presto card over the paper monthly pass.
Fare integration[edit]
GO Transit offers fare integration services with local transit providers, as well as with Via Rail.
Via Rail integration[edit]
Via Rail offers interline tickets with GO Transit, where passengers may travel on GO Transit trains and connect with Via Rail trains at any GO station also offering Via Rail services (Aldershot GO Station, Brampton GO Station, Georgetown GO Station, Guelph Central GO Station, Guildwood GO Station, Kitchener GO Station, Niagara Falls railway station (Ontario), Oakville GO Station, Oshawa GO Station, St. Catharines railway station, and Union Station) or vice versa on one ticket; the Via Rail ticket acts as a GO transit proof of payment on the GO transit leg. Via Rail interline tickets are not valid for GO buses when trains are not running.
Via Rail also offers the GO-Via Pak, which allows holders of GO Transit monthly passes (between applicable fare zones) to ride to Union Station from any GO Station also served by Via Rail trains (or vice versa) using Via Rail trains. The GO-Via Pak is priced at the difference between 10 adult single fares and the equivalent 10 adult economy fares on Via Rail. The GO-Via Pak is not offered for the seasonal trains serving St. Catharines railway station and Niagara Falls railway station (Ontario), or offered for trips originating at Guelph Central GO Station and Kitchener GO Station, due to Via Rail's own existing commuter pass options (note that Via Rail also offers commuter passes to Georgetown and Oshawa GO Stations, though they cannot be combined with GO Transit monthly passes in any event). The GO-Via Pak cannot be used with the Presto Card.
Local transit integration[edit]
Fare integration services may exist between GO Transit and local transit providers, where customers may ride GO buses within local service areas using appropriate local proof-of-payment (tickets and/or passes) at no additional cost; GO Transit will also issue a transfer for further travel on local transit providers upon request. Passengers paying by cash or travelling outside the local service area must pay GO transit fares. Passengers using Presto Card will be assessed GO transit fares, although plans are in place for local fares to be assessed instead of GO transit fares whenever they are lower.
Durham Region Transit - Though in the past, all GO bus routes serving Durham Region were integrated with DRT, since the launch of DRT Pulse only the routes not covered by DRT Pulse are integrated. This includes Route 71, 81, 88, 90, and 91, the latter two only between Newcastle and Downtown Oshawa.
York Region Transit - No routes currently are integrated with YRT, though GO Transit had operated routes both on behalf of YRT and as a complement to YRT service in the past, before its eventual replacement with YRT routes. The last such route to have been integrated was Route 69 Sutton, which was replaced by YRT Route 50 Queensville.
GO Transit co-fares[edit]
Local transit systems may offer reduced local fares, known as "co-fares", when making a connection between a local route and a GO train or bus at certain locations. In all cases, passengers must present GO Transit proof-of-payment (a monthly pass or an uncancelled ticket for passengers connecting to GO transit, or a monthly pass or a recently cancelled ticket for passengers connecting from GO Transit) to the local transit operator.
Passengers using the Presto Card will automatically be assessed the co-fare where appropriate, depending on the status of the local transit operator's integration with the Presto Card. If the local transit operator does not currently accept the Presto Card, the co-fare cannot be used with the Presto Card.
Transit Agency
Presto Card
Co-Fare
Co-Fare Monthly Pass
Notes
Barrie Transit No Free No Valid to/from Barrie South GO Station or Allandale Waterfront GO Station.
Despite Barrie Transit having not adopted the Presto Card on its buses, the co-fare is also accepted for Presto Card holders.
Brampton Transit
(includes Züm) Yes 80 cents No Valid to/from any GO train station in Brampton (Brampton, Bramalea, Mount Pleasant), Lisgar GO Station and Malton GO Station in Mississauga, and Brampton Bus Terminal.
Also valid for GO buses connecting to/from Brampton Transit services when using the Presto Card.
Co-fares are only accepted for single-ride tickets when connecting from GO Transit to BT services. A full BT fare is required when connecting to GO Transit from BT on a GO Transit single-ride ticket.
Burlington Transit Yes 70 cents No Valid to/from any GO train station in Burlington (Aldershot, Appleby, or Burlington).
Also valid for GO buses connecting to/from Burlington Transit services when using the Presto Card.
Durham Region Transit
(including DRT Pulse) Yes 75 cents No Valid to/from any GO Station in Durham Region (Ajax, Oshawa, Pickering, or Whitby) or Rouge Hill GO Station in Toronto.
Also valid for GO buses connecting to/from DRT services when using the Presto Card.
Co-fares are only accepted for single-ride tickets when connecting from GO Transit to DRT services. A full DRT fare is required when connecting to GO Transit from DRT on a GO Transit single-ride ticket.
Due to fare integration, DRT co-fares are not accepted on GO buses, and may not be used to connect between GO trains and GO buses; GO Transit fares will be assessed.
Grand River Transit
(including iXpress and GRT MobilityPLUS) No 50 cents No Valid to/from Kitchener GO Station.
Guelph Transit No 65 cents No Valid to/from Guelph Central GO Station and University of Guelph.
Co-fares are only accepted for single-ride tickets when connecting from GO Transit to Guelph Transit services. A full Guelph Transit fare is required when connected to GO Transit from Guelph Transit on a GO Transit single-ride ticket.
Milton Transit No 70 cents $8 Valid to/from Milton GO Station.
MiWay Yes 80 cents No Co-fares only offered for Presto Card holders.
Oakville Transit Yes 75 cents No Valid to/from any GO train station in Oakville (Bronte and Oakville), as well as Clarkson GO Station in Mississauga.
Also valid for GO buses connecting to/from Oakville Transit services when using the Presto Card.
Region of Peel TransHelp No 50 cents No Valid to/from Brampton GO Station.
York Region Transit
(includes Viva and Toronto Transit Commission routes serving York Region) Yes 75 cents No Valid to/from any GO train station in York Region (Aurora, Centennial, East Gwillimbury, King City, Langstaff, Lincolnville, Maple, Markham, Mount Joy, Newmarket, Richmond Hill, Rutherford, Stouffville, and Unionville), Milliken and York University GO Stations in Toronto, or Richmond Hill Centre Terminal.
Co-fares are only accepted for single-ride tickets when connecting from GO Transit to YRT services. A full YRT fare is required when connecting to GO Transit from YRT on a GO Transit single-ride ticket.
Enterprise Vivastation is considered part of Unionville GO Station for the purposes of transferring between GO Transit services and Viva Purple and Viva Green.
As the TTC does not currently implement the Presto Card on its buses, the Presto Card cannot be used with co-fares when connecting with TTC routes 102D Markham Road North (serving Mount Joy GO Station), 107B Dufferin North (serving Rutherford GO Station), 107D Keele North (serving Maple GO station), or 129A McCowan North (serving Centennial GO Station).
Though TTC Routes serving or terminating at Steeles Avenue may briefly enter York Region upon switching from outbound to inbound service, these are not services operated by the TTC on behalf of YRT; and thus co-fares are not offered. There are presently no TTC routes that serve both York Region and serve either of the GO Stations in Toronto where the YRT co-fare is offered.
The TTC Times Two program is not valid when connecting to a TTC bus in York Region, even if connecting to a destination in Toronto.
Co-Fare monthly passes are issued by the local transit agency, and allows for unlimited co-fare rides in a calendar month. Passengers not connecting to GO Transit services must pay the full local transit fare. Currently, co-fares are not counted towards any local transit agency's Presto card loyalty programs.
Other local transit integration[edit]
Other fare options are offered between GO Transit and local operators:
Hamilton Street Railway - Passengers using the Presto Card but without a HSR monthly pass loaded onto the card are entitled to a $1.60 discount when connecting to GO Transit from HSR and a $0.55 discount connecting to HSR from GO Transit.
Toronto Transit Commission - The TTC offers a program called "TTC Times Two", where TTC users may board a second TTC bus using a transfer from the first TTC leg if there is a GO Transit trip in between. Note that GO Transit proof of payment is not needed for this program. In addition, passengers connecting to TTC Route 38 (servicing University of Toronto's Scarborough campus) from Rouge Hill GO Station may travel for free with GO Transit proof of payment.
TTC Times Two is not offered for GO/TTC connections outside the City of Toronto, due to the fact that TTC operates routes on behalf of local transit agencies (MiWay and York Region Transit) when operating outside the City of Toronto.
As of January 7, 2018, Presto card holders who transfer from the GO Transit to the TTC (and vice-versa) can save up to $1.50 on their fares as part of a new co-fare discount agreement. It is not offered for customers who use cash, paper tickets, tokens, a Metropass or a monthly pass on Presto. [2]
References[edit]
Jump up ^ Kalinowski, Tess (15 October 2012). "GO Transit cancels monthly pass". Toronto Star. Retrieved 2012-10-15.
Jump up ^ https://www.prestocard.ca/en/about/discounted-double-fare
External links[edit]
GO Transit Fare Info
GO Transit Ticket Types
GO Transit Presto Card integration
GO Transit Fare Calculator
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