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Xenotransplantation

Xenotransplantation, or heterologous transplant, is the transplantation of living cells, tissues or organs from one species to another. Such cells, tissues or organs are called xenografts or xenotransplants. It is contrasted with allotransplantation, syngeneic transplantation or isotransplantation, and autotransplantation. Xenotransplantation is an artificial method of creating an animal-human chimera, that is, a human with a subset of animal cells. In contrast, an individual where each cell contains genetic material from a human and an animal is called a human–animal hybrid.

History
The first serious attempts at xenotransplantation (then called heterotransplantation) appeared in the scientific literature in 1905, when slices of rabbit kidney were transplanted into a child with chronic kidney disease. In the first two decades of the 20th century, several subsequent efforts to use organs from lambs, pigs, and primates were published. Starting in October 1963, doctors at Tulane University attempted renal transplantations from non-human primates in six people who were near death. The first person, a 32 year old woman with a chronic kidney disease received the kidneys of a rhesus monkey and the kidneys "functioned well for seven days, then failed," and the patient died later from her illness. The first successful attempt (one in which the patient was able to leave the hospital and return home) with a chimpanzee was performed on November 5 at Charity Hospital in New Orleans by a 12-man team of Tulane physicians, led by Keith Reemtsma, and the patient, a 44-year-old dock worker named Jefferson Davis, left the hospital on December 17 after a six-week recuperation. After this and several subsequent unsuccessful attempts to use primates as organ donors and the development of a working cadaver organ procuring program, interest in xenotransplantation for kidney failure dissipated. The procedure was performed by Leonard Lee Bailey at Loma Linda University Medical Center in Loma Linda, California. Fae died 21 days later, on November 15, due to a humoral-based graft rejection thought to be caused mainly by an ABO blood type mismatch, considered unavoidable due to the rarity of type O baboons. The graft was meant to be temporary, but unfortunately a suitable allograft replacement could not be found in time. While the procedure itself did not advance the progress on xenotransplantation, it did shed a light on the insufficient amount of organs for infants. The story made such an impact that the crisis of infant organ shortage improved for that time. Non-human heart, lungs, and kidneys to a human The first heart transplant in a human ever performed was by Hardy in 1964, using a chimpanzee heart, but the patient died within 2 hours. pig's heart, lungs and kidneys into a human was performed in Sonapur, Assam in mid-December 1996, and was announced in January 1997. The Indian cardiothoracic surgeon Dhani Ram Baruah and two of his associates, Jonathan Ho Kei-shing (of the Hong Kong-based Prince of Wales Medical Institute) and C.S. James, performed the surgeries. All three involved in the surgery were arrested on January 9, 1997, Baruah was dismissed in medical circles as a "mad scientist" and the procedure was dubbed a "hoax". Baruah signed a statement saying he had done no transplant, but then alleged that the confession was forced from him. Dhani Ram Baruah's surgical institute was also found to be without necessary registration. Critics said Dhani Bam Baruah's claims and medical procedures were neither taken seriously nor accepted by the scientific community because he never got his findings scientifically peer-reviewed. Past complaints of ethics violations during surgeries in Hong Kong by Baruah and Ho had occurred in 1992, when they had implanted heart valves, developed by Baruah, made of animal tissue. A year later, six patients died. The Asian Medical News reported that "grave concerns" were expressed "over the procedure and ethics of the implementation". The kidney was procured from a pig with only a single gene modification: the removal of alpha-gal. In July 2023, surgeons from the NYU Langone Transplant Institute completed a transplant of a genetically modified pig kidney (along with the pig's thymus gland underneath it) into a patient declared brain dead but maintained on a respirator. The patient had previously consented to be an organ donor, but his tissues were not considered suitable for transplant. The kidney came from an animal with a knocked-out gene for the production of alpha gal sugars, which has been implicated in immune response to mammalian tissue. In order to ensure that renal function was only supported by the pig kidney, the team removed both of the patient's kidneys. The team has reported that the kidney has maintained optimal functioning for over a month, as evidenced by routine testing of creatinine and weekly biopsies. The team plans to monitor the patient for another month, pending approval by ethics board and his family. In March 2024, Richard Slayman, a patient whose transplanted human kidney had failed, received a genetically engineered pig kidney xenotransplant from surgeons at Massachusetts General Hospital. This kidney has 69 genomic edits (3 gene knockouts, 7 human gene insertions and 59 copies of the porcine retrovirus knockout) made by eGenesis, Inc. Slayman died a few months later of unrelated causes, with no apparent rejection of the kidney. Meanwhile, in April 2024, Lisa Pisano became the second person to receive such a kidney transplant. Because of "unique challenges" related to a mechanical heart pump she received along with the kidney, her kidney had to be removed due to "insufficient blood flow" late in May. Medication also deteriorated the kidney, which led to its rejection. In 2025, Tim Andrews and Bill Stewart received gene-edited pig kidneys from surgeons at Massachusetts General Hospital. Andrews lived with the kidney for 271 days, doubling the previous record of 130 days for a pig kidney transplant, but just short of the xenotransplant record of 273 days for a chimpanzee kidney transplant in 1964. The kidney was removed in October 2025 due to rejection, and Andrews later received a human kidney transplant in January 2026. "Based on lessons from these early cases, the U.S. Food and Drug Administration (FDA) has granted approval to biotech company eGenesis to launch a clinical trial that will transplant gene-edited pig kidneys into 30 patients who are 50 or older and on dialysis while awaiting a human kidney." at the University of Maryland Medical Center and University of Maryland School of Medicine performed a heart transplant from a genetically modified pig to a terminally ill patient, David Bennett Sr., who was ineligible for a standard human heart transplant. The pig had undergone specific gene editing to remove enzymes responsible for producing sugar antigens that lead to hyperacute organ rejection in humans. The US medical regulator gave special dispensation to carry out the procedure under compassionate use criteria. The recipient died two months after the transplantation. In June and July 2022, surgeons at NYU Langone Health performed two genetically modified pig heart transplants into recently deceased humans. The hearts were from pigs that had the identical 10 genetic modifications used in the University of Maryland Medical Center heart xenotransplantation in January 2022. All three hearts came from Revivicor, Inc., a facility based in Blacksburg, Va., and a subsidiary of United Therapeutics. On 20 September 2023, surgeons at the University of Maryland Medical Center in Baltimore performed a heart transplant from a genetically modified pig to Lawrence Faucette, a patient with terminal heart disease who was ineligible for a traditional heart transplant. On 30 October 2023, Faucette died after showing signs of organ rejection. ==Potential uses==
Potential uses
A worldwide shortage of organs for clinical implantation causes about 20–35% of patients who need replacement organs to die on the waiting list. Certain procedures, some of which are being investigated in early clinical trials, aim to use cells or tissues from other species to treat life-threatening and debilitating illnesses such as cancer, diabetes, liver failure and Parkinson's disease. If vitrification can be perfected, it could allow for long-term storage of xenogenic cells, tissues and organs so that they would be more readily available for transplant. Xenotransplants could save 100,000 patients waiting for donated organs. Xenotransplantation of human tumor cells into immunocompromised mice is a research technique frequently used in oncology research. It is used to predict the sensitivity of the transplanted tumor to various cancer treatments; several companies offer this service, including the Jackson Laboratory. Human organs have been transplanted into animals as a powerful research technique for studying human biology without harming human patients. This technique has also been proposed as an alternative source of human organs for future transplantation into human patients. For example, researchers from the Ganogen Research Institute transplanted human fetal kidneys into rats which demonstrated life supporting function and growth. == Potential animal organ donors ==
Potential animal organ donors
Since they are the closest relatives to humans, non-human primates were first considered as a potential organ source for xenotransplantation to humans. Chimpanzees were originally considered the best option since their organs are of similar size, and they have good blood type compatibility with humans, which makes them potential candidates for xenotransfusions. However, since chimpanzees are listed as an endangered species, other potential donors were sought. Baboons are more readily available, but impractical as potential donors. Problems include their smaller body size, the infrequency of blood group O (the universal donor), their long gestation period, and their typically small number of offspring. In addition, a major problem with the use of nonhuman primates is the increased risk of disease transmission, since they are so closely related to humans. Pigs (Sus scrofa domesticus) are currently thought to be the best candidates for organ donation. The risk of cross-species disease transmission is decreased because of their increased phylogenetic distance from humans. They are inexpensive and easy to maintain in pathogen-free facilities, and current gene editing tools are adapted to pigs to combat rejection and potential zoonoses. Treatments sourced from pigs have proven to be successful such as porcine-derived insulin for patients with diabetes mellitus. Increasingly, genetically engineered pigs are becoming the norm, which raises moral qualms, but also increases the success rate of the transplant. Current experiments in xenotransplantation most often use pigs as the donor, and baboons as human models. In 2020, the U.S. Food and Drug Administration approved a genetic modification of pigs so they do not produce alpha-gal sugars. Pig organs have been used for kidney and heart transplants into humans. ==Barriers and issues==
Barriers and issues
Immunologic barriers As of 20 September 2025, no xenotransplantation trials have been successful for more than 238 days due to the many obstacles arising from the response of the recipient's immune system, although at least two patients have not yet rejected their xenotransplants and are still off dialysis. This response, which is generally more extreme than in allotransplantations, ultimately results in rejection of the xenograft, and can in some cases result in the immediate death of the recipient. There are several types of rejection organ xenografts are faced with, these include hyperacute rejection, acute vascular rejection, cellular rejection, and chronic rejection. A rapid, violent, and hyperacute response comes as a result of antibodies present in the host organism. These antibodies are known as xenoreactive natural antibodies (XNAs). Most non-primates contain this enzyme thus, this epitope is present on the organ epithelium and is perceived as a foreign antigen by primates, which lack the galactosyl transferase enzyme. In pig to primate xenotransplantation, XNAs recognize porcine glycoproteins of the integrin family. Overcoming hyperacute rejection Since hyperacute rejection presents such a barrier to the success of xenografts, several strategies to overcome it are under investigation: Interruption of the complement cascade • The recipient's complement cascade can be inhibited through the use of cobra venom factor (which depletes C3), soluble complement receptor type 1, anti-C5 antibodies, or C1 inhibitor (C1-INH). Disadvantages of this approach include the toxicity of cobra venom factor, and most importantly these treatments would deprive the individual of a functional complement system. • Increased expression of H-transferase (α-1,2-fucosyltransferase), an enzyme that competes with galactosyl transferase. Experiments have shown this reduces α-Gal expression by 70%. • Expression of human complement regulators (CD55, CD46, and CD59) to inhibit the complement cascade. • Plasmaphoresis, on humans to remove 1,3 galactosyltransferase, reduces the risk of activation of effector cells such as CTL (CD8 T cells), complement pathway activation and delayed type hypersensitivity (DTH). Acute vascular rejection Also known as delayed xenoactive rejection, this type of rejection occurs in discordant xenografts within 2 to 3 days, if hyperacute rejection is prevented. The process is much more complex than hyperacute rejection and is currently not completely understood. Acute vascular rejection requires de novo protein synthesis and is driven by interactions between the graft endothelial cells and host antibodies, macrophages, and platelets. The response is characterized by an inflammatory infiltrate of mostly macrophages and natural killer cells (with small numbers of T cells), intravascular thrombosis, and fibrinoid necrosis of vessel walls. when the complement cascade is interrupted, circulating antibodies are removed, their function is changed, or there is a change in the expression of surface antigens on the graft. This allows the xenograft to up-regulate and express protective genes, which aid in resistance to injury, such as heme oxygenase-1 (an enzyme that catalyzes the degradation of heme). The strength of cellular rejection in xenografts remains uncertain, however, it is expected to be stronger than in allografts due to differences in peptides among different animals. This leads to more antigens potentially recognized as foreign, thus eliciting a greater indirect xenogenic response. Donor stem cells are introduced into the bone marrow of the recipient, where they coexist with the recipient's stem cells. The bone marrow stem cells give rise to cells of all hematopoietic lineages, through the process of hematopoiesis. Lymphoid progenitor cells are created by this process and move to the thymus where negative selection eliminates T cells found to be reactive to self. The existence of donor stem cells in the recipient's bone marrow causes donor reactive T cells to be considered self-reactive and undergo apoptosis. Dysregulated coagulation Successful efforts have been made to create knockout mice without α1,3GT; the resulting reduction in the highly immunogenic αGal epitope has resulted in the reduction of the occurrence of hyperacute rejection, but has not eliminated other barriers to xenotransplantation such as dysregulated coagulation, also known as coagulopathy. Different organ xenotransplants result in different responses in clotting. For example, kidney transplants result in a higher degree of coagulopathy, or impaired clotting, than cardiac transplants, whereas liver xenografts result in severe thrombocytopenia, causing recipient death within a few days due to bleeding. Additionally, spontaneous platelet accumulation may be caused by contact with pig von Willebrand factor. Xenotransplantation may increase the chance of disease transmission for 3 reasons: (1) implantation breaches the physical barrier that normally helps to prevent disease transmission, (2) the recipient of the transplant will be severely immunosuppressed, and (3) human complement regulators (CD46, CD55, and CD59) expressed in transgenic pigs have been shown to serve as virus receptors, and may also help to protect viruses from attack by the complement system. Examples of viruses carried by pigs include porcine herpesvirus, rotavirus, parvovirus, and circovirus. Porcine herpesviruses and rotaviruses can be eliminated from the donor pool by screening, however others (such as parvovirus and circovirus) may contaminate food and footwear then re-infect the herd. Thus, pigs to be used as organ donors must be housed under strict regulations and screened regularly for microbes and pathogens. Unknown viruses, as well as those not harmful in the animal, may also pose risks. Since then PERVs have been eliminated from the pig genomes in successfully donated pig kidneys. This was repealed in 2009 after an NHMRC review stated "... the risks, if appropriately regulated, are minimal and acceptable given the potential benefits.", citing international developments on the management and regulation of xenotransplantation by the World Health Organisation and the European Medicines Agency. Porcine endogenous retroviruses Endogenous retroviruses are remnants of ancient viral infections, found in the genomes of most, if not all, mammalian species. Integrated into the chromosomal DNA, they are vertically transferred through inheritance. Most breeds of swine harbor approximately 50 PERV genomes in their DNA. Although it is likely that most of these are defective, some may be able to produce infectious viruses so every proviral genome must be sequenced to identify which ones pose a threat. In addition, through complementation and genetic recombination, two defective PERV genomes could give rise to an infectious virus. There are three subgroups of infectious PERVs (PERV-A, PERV-B and PERV-C). Experiments have shown that PERV-A and PERV-B can infect human cells in culture. To date no experimental xenotransplantations have demonstrated PERV transmission, yet this does not mean PERV infections in humans are impossible. and eliminated infection from the pig to human cells in culture. Ethics Xenografts have been a controversial procedure since they were first attempted. Many, including animal rights groups, strongly oppose killing animals to harvest their organs for human use. In the 1960s, many organs came from the chimpanzees, and were transferred into people that were deathly ill, and in turn, did not live much longer afterwards. Modern scientific supporters of xenotransplantation argue that the potential benefits to society outweigh the risks, making pursuing xenotransplantation the moral choice. None of the major religions object to the use of genetically modified pig organs for life-saving transplantation. Religions such as Buddhism and Jainism, however, have long espoused non-violence towards all living creatures. History of xenotransplantation in ethics At the beginning of the 20th century when studies in xenotransplantation were just beginning, few questioned the morality of it, turning to animals as a "natural" alternative to allografts. While satirical plays mocked xenografters such as Serge Voronoff, and some images showing emotionally distraught primates – whom Voronoff had deprived of their testicles – appeared, no serious attempts were yet made to question the science based on animal rights concerns. Despite this, it is considerably unlikely that animal suffering will provide sufficient impetus for regulators to prevent xenotransplantation. The patient should understand the risks and benefits of such a transplantation. A public health dimension can also be considered. The Ethics Committee of the International Xenotransplantation Association pointed out in 2003 that one major ethical issue is the societal response to such a procedure. The application of the four bioethics principles is standardized in the moral conduct of laboratories. The four principles emphasize informed consent, the Hippocratic Oath to do no harm, using skills to help others, and protecting the right to quality care. Though xenotransplantation may have future medical benefits, it also has the serious risk of introducing and spreading the infectious diseases, into the human population. Guidelines have been drafted by governments with the purpose of forming the foundation of infectious disease surveillance. ==See also==
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