Symbiogenesis

From Wikipedia, the free encyclopedia

https://en.wikipedia.org/wiki/Symbiogenesis 

In the theory of symbiogenesis, a merger of an archaean and an aerobic bacterium created the eukaryotes, with aerobic mitochondria; a second merger added chloroplasts, creating the green plants. The original theory by Lynn Margulis proposed an additional preliminary merger, but this is poorly supported and not now generally believed.

Symbiogenesis (endosymbiotic theory, or serial endosymbiotic theory) is the leading evolutionary theory of the origin of eukaryotic cells from prokaryotic organisms. The theory holds that mitochondria, plastids such as chloroplasts, and possibly other organelles of eukaryotic cells are descended from formerly free-living prokaryotes (more closely related to the Bacteria than to the Archaea) taken one inside the other in endosymbiosis. Mitochondria appear to be phylogenetically related to Rickettsiales bacteria, while chloroplasts are thought to be related to cyanobacteria.

The idea that chloroplasts were originally independent organisms that merged into a symbiotic relationship with other one-celled organisms dates back to the 19th century, when it was espoused by researchers such as Andreas Schimper. The endosymbiotic theory was articulated in 1905 and 1910 by the Russian botanist Konstantin Mereschkowski, and advanced and substantiated with microbiological evidence by Lynn Margulis in 1967.

Among the many lines of evidence supporting symbiogenesis are that mitochondria and plastids contain their own chromosomes and reproduce by splitting in two, parallel but separate from the sexual reproduction of the rest of the cell; that the chromosomes of some mitochondria and plastids are single circular DNA molecules similar to the circular chromosomes of bacteria; that the transport proteins called porins are found in the outer membranes of mitochondria and chloroplasts, and also bacterial cell membranes; and that cardiolipin is found only in the inner mitochondrial membrane and bacterial cell membranes.

History

Konstantin Mereschkowski's 1905 tree-of-life diagram, showing the origin of complex life-forms by two episodes of symbiogenesis, the incorporation of symbiotic bacteria to form successively nuclei and chloroplasts

The Russian botanist Konstantin Mereschkowski first outlined the theory of symbiogenesis (from Greek: σύν syn "together", βίος bios "life", and γένεσις genesis "origin, birth") in his 1905 work, The nature and origins of chromatophores in the plant kingdom, and then elaborated it in his 1910 The Theory of Two Plasms as the Basis of Symbiogenesis, a New Study of the Origins of Organisms. Mereschkowski proposed that complex life-forms had originated by two episodes of symbiogenesis, the incorporation of symbiotic bacteria to form successively nuclei and chloroplasts. Mereschkowski knew of the work of botanist Andreas Schimper. In 1883, Schimper had observed that the division of chloroplasts in green plants closely resembled that of free-living cyanobacteria. Schimper had tentatively proposed (in a footnote) that green plants had arisen from a symbiotic union of two organisms. In 1918 the French scientist Paul Jules Portier published Les Symbiotes, in which he claimed that the mitochondria originated from a symbiosis process. Ivan Wallin advocated the idea of an endosymbiotic origin of mitochondria in the 1920s. The Russian botanist Boris Kozo-Polyansky became the first to explain the theory in terms of Darwinian evolution. In his 1924 book A New Principle of Biology. Essay on the Theory of Symbiogenesis, he wrote, "The theory of symbiogenesis is a theory of selection relying on the phenomenon of symbiosis."

These theories did not gain traction until more detailed electron-microscopic comparisons between cyanobacteria and chloroplasts were made, such as by Hans Ris in 1961 and 1962. These, combined with the discovery that plastids and mitochondria contain their own DNA, led to a resurrection of the idea of symbiogenesis in the 1960s. Lynn Margulis advanced and substantiated the theory with microbiological evidence in a 1967 paper, On the origin of mitosing cells. In her 1981 work Symbiosis in Cell Evolution she argued that eukaryotic cells originated as communities of interacting entities, including endosymbiotic spirochaetes that developed into eukaryotic flagella and cilia. This last idea has not received much acceptance, because flagella lack DNA and do not show ultrastructural similarities to bacteria or to archaea (see also: Evolution of flagella and Prokaryotic cytoskeleton). According to Margulis and Dorion Sagan, "Life did not take over the globe by combat, but by networking" (i.e., by cooperation). Christian de Duve proposed that the peroxisomes may have been the first endosymbionts, allowing cells to withstand growing amounts of free molecular oxygen in the Earth's atmosphere. However, it now appears that peroxisomes may be formed de novo, contradicting the idea that they have a symbiotic origin. The fundamental theory of symbiogenesis as the origin of mitochondria and chloroplasts is now widely accepted.

Symbiogenesis revolutionized the history of evolution by proposing a mechanism for evolutionary development not encompassed in the original Darwininan vision. Symbiogenesis demonstrated that major evolutionary advancements, particularly the origin of eukaryotic cells, may have resulted from symbiotic mergers rather than from gradual mutations and individual competition, i.e., classical natural selection. Accordingly, symbiogenic theory suggests that endosymbiosis may be a powerful force in generating evolutionary novelty, beyond that which can be explained by natural selection alone.

From endosymbionts to organelles

An autogenous model of the origin of eukaryotic cells. Evidence now shows that a mitochondrion-less eukaryote has never existed, i.e. the nucleus was acquired at the same time as the mitochondria.

Biologists usually distinguish organelles from endosymbionts – whole organisms living inside other organisms – by their reduced genome sizes. As an endosymbiont evolves into an organelle, most of its genes are transferred to the host cell genome. The host cell and organelle therefore need to develop a transport mechanism that enables the return of the protein products needed by the organelle but now manufactured by the cell.

Free-living ancestors

Alphaproteobacteria were formerly thought to be the free-living organisms most closely related to mitochondria. Later research indicates that mitochondria are most closely related to Pelagibacterales bacteria, in particular, those in the SAR11 clade.

Nitrogen-fixing filamentous cyanobacteria are the free-living organisms most closely related to plastids.

Both cyanobacteria and alphaproteobacteria maintain a large (>6 Mb) genome encoding thousands of proteins. Plastids and mitochondria exhibit a dramatic reduction in genome size when compared with their bacterial relatives. Chloroplast genomes in photosynthetic organisms are normally 120–200 kb encoding 20–200 proteins and mitochondrial genomes in humans are approximately 16 kb and encode 37 genes, 13 of which are proteins. Using the example of the freshwater amoeboid, however, Paulinella chromatophora, which contains chromatophores found to be evolved from cyanobacteria, Keeling and Archibald argue that this is not the only possible criterion; another is that the host cell has assumed control of the regulation of the former endosymbiont's division, thereby synchronizing it with the cell's own division. Nowack and her colleagues gene sequenced the chromatophore (1.02 Mb) and found that only 867 proteins were encoded by these photosynthetic cells. Comparisons with their closest free living cyanobacteria of the genus Synechococcus (having a genome size 3 Mb, with 3300 genes) revealed that chromatophores had undergone a drastic genome shrinkage. Chromatophores contained genes that were accountable for photosynthesis but were deficient in genes that could carry out other biosynthetic functions; this observation suggests that these endosymbiotic cells are highly dependent on their hosts for their survival and growth mechanisms. Thus, these chromatophores were found to be non-functional for organelle-specific purposes when compared with mitochondria and plastids. This distinction could have promoted the early evolution of photosynthetic organelles.

The loss of genetic autonomy, that is, the loss of many genes from endosymbionts, occurred very early in evolutionary time. Taking into account the entire original endosymbiont genome, there are three main possible fates for genes over evolutionary time. The first is the loss of functionally redundant genes, in which genes that are already represented in the nucleus are eventually lost. The second is the transfer of genes to the nucleus, while the third is that genes remain in the organelle that was once an organism. The loss of autonomy and integration of the endosymbiont with its host can be primarily attributed to nuclear gene transfer. As organelle genomes have been greatly reduced over evolutionary time, nuclear genes have expanded and become more complex. As a result, many plastid and mitochondrial processes are driven by nuclear encoded gene products. In addition, many nuclear genes originating from endosymbionts have acquired novel functions unrelated to their organelles.

Gene transfer mechanisms

The mechanisms of gene transfer are not fully known; however, multiple hypotheses exist to explain this phenomenon. The possible mechanisms include the Complementary DNA (cDNA) hypothesis and the bulk flow hypothesis.

The cDNA hypothesis involves the use of messenger RNA (mRNAs) to transport genes from organelles to the nucleus where they are converted to cDNA and incorporated into the genome. The cDNA hypothesis is based on studies of the genomes of flowering plants. Protein coding RNAs in mitochondria are spliced and edited using organelle-specific splice and editing sites. Nuclear copies of some mitochondrial genes, however, do not contain organelle-specific splice sites, suggesting a processed mRNA intermediate. The cDNA hypothesis has since been revised as edited mitochondrial cDNAs are unlikely to recombine with the nuclear genome and are more likely to recombine with their native mitochondrial genome. If the edited mitochondrial sequence recombines with the mitochondrial genome, mitochondrial splice sites would no longer exist in the mitochondrial genome. Any subsequent nuclear gene transfer would therefore also lack mitochondrial splice sites.

The bulk flow hypothesis is the alternative to the cDNA hypothesis, stating that escaped DNA, rather than mRNA, is the mechanism of gene transfer. According to this hypothesis, disturbances to organelles, including autophagy (normal cell destruction), gametogenesis (the formation of gametes), and cell stress release DNA which is imported into the nucleus and incorporated into the nuclear DNA using non-homologous end joining (repair of double stranded breaks). For example, in the initial stages of endosymbiosis, due to a lack of major gene transfer, the host cell had little to no control over the endosymbiont. The endosymbiont underwent cell division independently of the host cell, resulting in many "copies" of the endosymbiont within the host cell. Some of the endosymbionts lysed (burst), and high levels of DNA were incorporated into the nucleus. A similar mechanism is thought to occur in tobacco plants, which show a high rate of gene transfer and whose cells contain multiple chloroplasts. In addition, the bulk flow hypothesis is also supported by the presence of non-random clusters of organelle genes, suggesting the simultaneous movement of multiple genes.

Ford Doolittle proposed that (whatever the mechanism) gene transfer behaves like a ratchet, resulting in unidirectional transfer of genes from the organelle to the nuclear genome. When genetic material from an organelle is incorporated into the nuclear genome, either the organelle or nuclear copy of the gene may be lost from the population. If the organelle copy is lost and this is fixed, or lost through genetic drift, a gene is successfully transferred to the nucleus. If the nuclear copy is lost, horizontal gene transfer can occur again, and the cell can 'try again' to have successful transfer of genes to the nucleus. In this ratchet-like way, genes from an organelle would be expected to accumulate in the nuclear genome over evolutionary time.

Endosymbiosis of protomitochondria

Further information: Eukaryogenesis

Endosymbiotic theory for the origin of mitochondria suggests that the proto-eukaryote engulfed a protomitochondrion, and this endosymbiont became an organelle, a major step in eukaryogenesis, the creation of the eukaryotes.

Mitochondria

Internal symbiont: mitochondrion has a matrix and membranes, like a free-living alphaproteobacterial cell, from which it may derive.

Mitochondria are organelles that synthesize the energy-carrying molecule ATP for the cell by metabolizing carbon-based macromolecules. The presence of DNA in mitochondria and proteins, derived from mtDNA, suggest that this organelle may have been a prokaryote prior to its integration into the proto-eukaryote. Mitochondria are regarded as organelles rather than endosymbionts because mitochondria and the host cells share some parts of their genome, undergo division simultaneously, and provide each other with means to produce energy. The endomembrane system and nuclear membrane were hypothesized to have derived from the protomitochondria.

Nuclear membrane

The presence of a nucleus is one major difference between eukaryotes and prokaryotes. Some conserved nuclear proteins between eukaryotes and prokaryotes suggest that these two types had a common ancestor. Another theory behind nucleation is that early nuclear membrane proteins caused the cell membrane to fold and form a sphere with pores like the nuclear envelope. As a way of forming a nuclear membrane, endosymbiosis could be expected to use less energy than if the cell was to develop a metabolic process to fold the cell membrane for the purpose. Digesting engulfed cells without energy-producing mitochondria would have been challenging for the host cell. On this view, membrane-bound bubbles or vesicles leaving the protomitochondria may have formed the nuclear envelope.

The process of symbiogenesis by which the early eukaryotic cell integrated the proto-mitochondrion likely included protection of the archaeal host genome from the release of reactive oxygen species. These would have been formed during oxidative phosphorylation and ATP production by the proto-mitochondrion. The nuclear membrane may have evolved as an adaptive innovation for protecting against nuclear genome DNA damage caused by reactive oxygen species. Substantial transfer of genes from the ancestral proto-mitochondrial genome to the nuclear genome likely occurred during early eukaryotic evolution. The greater protection of the nuclear genome against reactive oxygen species afforded by the nuclear membrane may explain the adaptive benefit of this gene transfer.

Endomembrane system

Diagram of endomembrane system in eukaryotic cell

Modern eukaryotic cells use the endomembrane system to transport products and wastes in, within, and out of cells. The membrane of nuclear envelope and endomembrane vesicles are composed of similar membrane proteins. These vesicles also share similar membrane proteins with the organelle they originated from or are traveling towards. This suggests that what formed the nuclear membrane also formed the endomembrane system. Prokaryotes do not have a complex internal membrane network like eukaryotes, but they could produce extracellular vesicles from their outer membrane. After the early prokaryote was consumed by a proto-eukaryote, the prokaryote would have continued to produce vesicles that accumulated within the cell. Interaction of internal components of vesicles may have led to the endoplasmic reticulum and the Golgi apparatus, both being parts of the endomembrane system.

Cytoplasm

The syntrophy hypothesis, proposed by López-García and Moreira in 1998, suggested that eukaryotes arose by combining the metabolic capabilities of an archaean, a fermenting deltaproteobacterium, and a methanotrophic alphaproteobacterium which became the mitochondrion. In 2020, the same team updated their syntrophy proposal to cover an promethearchaeon that produced hydrogen with deltaproteobacterium that oxidised sulphur. A third organism, an alphaproteobacterium able to respire both aerobically and anaerobically, and to oxidise sulphur, developed into the mitochondrion; it may possibly also have been able to photosynthesise.

Date

The question of when the transition from prokaryotic to eukaryotic form occurred and when the first crown group eukaryotes appeared on earth is unresolved. The oldest known body fossils that can be positively assigned to the Eukaryota are acanthomorphic acritarchs from the 1.631 Gya Deonar Formation of India. These fossils can still be identified as derived post-nuclear eukaryotes with a sophisticated, morphology-generating cytoskeleton sustained by mitochondria. This fossil evidence indicates that endosymbiotic acquisition of alphaproteobacteria must have occurred before 1.6 Gya. Molecular clocks have also been used to estimate the last eukaryotic common ancestor, however these methods have large inherent uncertainty and give a wide range of dates. Reasonable results include the estimate of c. 1.8 Gya. A 2.3 Gya estimate also seems reasonable, and has the added attraction of coinciding with one of the most pronounced biogeochemical perturbations in Earth history, the early Palaeoproterozoic Great Oxygenation Event. The marked increase in atmospheric oxygen concentrations at that time has been suggested as a contributing cause of eukaryogenesis, inducing the evolution of oxygen-detoxifying mitochondria. Alternatively, the Great Oxidation Event might be a consequence of eukaryogenesis, and its impact on the export and burial of organic carbon.

Organellar genomes

Plastomes and mitogenomes

The human mitochondrial genome has retained genes encoding 2 rRNAs (blue), 22 tRNAs (white), and 13 redox proteins (yellow, orange, red).

Some endosymbiont genes remain in the organelles. Plastids and mitochondria retain genes encoding rRNAs, tRNAs, proteins involved in redox reactions, and proteins required for transcription, translation, and replication. There are many hypotheses to explain why organelles retain a small portion of their genome; however no one hypothesis will apply to all organisms, and the topic is still quite controversial. The hydrophobicity hypothesis states that highly hydrophobic (water hating) proteins (such as the membrane bound proteins involved in redox reactions) are not easily transported through the cytosol and therefore these proteins must be encoded in their respective organelles. The code disparity hypothesis states that the limit on transfer is due to differing genetic codes and RNA editing between the organelle and the nucleus. The redox control hypothesis states that genes encoding redox reaction proteins are retained in order to effectively couple the need for repair and the synthesis of these proteins. For example, if one of the photosystems is lost from the plastid, the intermediate electron carriers may lose or gain too many electrons, signalling the need for repair of a photosystem. The time delay involved in signalling the nucleus and transporting a cytosolic protein to the organelle results in the production of damaging reactive oxygen species. The final hypothesis states that the assembly of membrane proteins, particularly those involved in redox reactions, requires coordinated synthesis and assembly of subunits; however, translation and protein transport coordination is more difficult to control in the cytoplasm.

Non-photosynthetic plastid genomes

The majority of the genes in the mitochondria and plastids are related to the expression (transcription, translation and replication) of genes encoding proteins involved in either photosynthesis (in plastids) or cellular respiration (in mitochondria). One might predict that the loss of photosynthesis or cellular respiration would allow for the complete loss of the plastid genome or the mitochondrial genome respectively. While there are numerous examples of mitochondrial descendants (mitosomes and hydrogenosomes) that have lost their entire organellar genome, non-photosynthetic plastids tend to retain a small genome. There are two main hypotheses to explain this occurrence:

The essential tRNA hypothesis notes that there have been no documented functional plastid-to-nucleus gene transfers of genes encoding RNA products (tRNAs and rRNAs). As a result, plastids must make their own functional RNAs or import nuclear counterparts. The genes encoding tRNA-Glu and tRNA-fmet, however, appear to be indispensable. The plastid is responsible for haem biosynthesis, which requires plastid encoded tRNA-Glu (from the gene trnE) as a precursor molecule. Like other genes encoding RNAs, trnE cannot be transferred to the nucleus. In addition, it is unlikely trnE could be replaced by a cytosolic tRNA-Glu as trnE is highly conserved; single base changes in trnE have resulted in the loss of haem synthesis. The gene for tRNA-formylmethionine (tRNA-fmet) is also encoded in the plastid genome and is required for translation initiation in both plastids and mitochondria. A plastid is required to continue expressing the gene for tRNA-fmet so long as the mitochondrion is translating proteins.

The limited window hypothesis offers a more general explanation for the retention of genes in non-photosynthetic plastids. According to this hypothesis, genes are transferred to the nucleus following the disturbance of organelles. Disturbance was common in the early stages of endosymbiosis, however, once the host cell gained control of organelle division, eukaryotes could evolve to have only one plastid per cell. Having only one plastid severely limits gene transfer as the lysis of the single plastid would likely result in cell death. Consistent with this hypothesis, organisms with multiple plastids show an 80-fold increase in plastid-to-nucleus gene transfer compared with organisms with single plastids.

Evidence

There are many lines of evidence that mitochondria and plastids including chloroplasts arose from bacteria.

• New mitochondria and plastids are formed only through binary fission, the form of cell division used by bacteria and archaea.
• If a cell's mitochondria or chloroplasts are removed, the cell does not have the means to create new ones. In some algae, such as Euglena, the plastids can be destroyed by certain chemicals or prolonged absence of light without otherwise affecting the cell: the plastids do not regenerate.
• Transport proteins called porins are found in the outer membranes of mitochondria and chloroplasts and are also found in bacterial cell membranes.
• A membrane lipid cardiolipin is exclusively found in the inner mitochondrial membrane and bacterial cell membranes.
• Some mitochondria and some plastids contain single circular DNA molecules that are similar to the DNA of bacteria both in size and structure.
• Genome comparisons suggest a close relationship between mitochondria and Alphaproteobacteria.
• Genome comparisons suggest a close relationship between plastids and cyanobacteria.
• Many genes in the genomes of mitochondria and chloroplasts have been lost or transferred to the nucleus of the host cell. Consequently, the chromosomes of many eukaryotes contain genes that originated from the genomes of mitochondria and plastids.
• Mitochondria and plastids contain their own ribosomes; these are more similar to those of bacteria (70S) than those of eukaryotes.
• Proteins created by mitochondria and chloroplasts use N-formylmethionine as the initiating amino acid, as do proteins created by bacteria but not proteins created by eukaryotic nuclear genes or archaea.

Comparison of chloroplasts and cyanobacteria showing their similarities. Both chloroplasts and cyanobacteria have a double membrane, DNA, ribosomes, and chlorophyll-containing thylakoids.

Secondary endosymbiosis

See also: Plastid evolution § Secondary endosymbiosis

Primary endosymbiosis involves the engulfment of a cell by another free living organism. Secondary endosymbiosis occurs when the product of primary endosymbiosis is itself engulfed and retained by another free living eukaryote. Secondary endosymbiosis has occurred several times and has given rise to extremely diverse groups of algae and other eukaryotes. Some organisms can take opportunistic advantage of a similar process, where they engulf an alga and use the products of its photosynthesis, but once the prey item dies (or is lost) the host returns to a free living state. Obligate secondary endosymbionts become dependent on their organelles and are unable to survive in their absence. A secondary endosymbiosis event involving an ancestral red alga and a heterotrophic eukaryote resulted in the evolution and diversification of several other photosynthetic lineages including Cryptophyta, Haptophyta, Stramenopiles (or Heterokontophyta), and Alveolata.

A possible secondary endosymbiosis has been observed in process in the heterotrophic protist Hatena. This organism behaves like a predator until it ingests a green alga, which loses its flagella and cytoskeleton but continues to live as a symbiont. Hatena meanwhile, now a host, switches to photosynthetic nutrition, gains the ability to move towards light, and loses its feeding apparatus.

Despite the diversity of organisms containing plastids, the morphology, biochemistry, genomic organisation, and molecular phylogeny of plastid RNAs and proteins suggest a single origin of all extant plastids – although this theory was still being debated in 2008.

Nitroplasts

A unicellular marine alga, Braarudosphaera bigelowii (a coccolithophore, which is a eukaryote), has been found with a cyanobacterium as an endosymbiont. The cyanobacterium forms a nitrogen-fixing structure, dubbed the nitroplast. It divides evenly when the host cell undergoes mitosis, and many of its proteins derive from the host alga, implying that the endosymbiont has proceeded far along the path towards becoming an organelle. The cyanobacterium is named Candidatus Atelocyanobacterium thalassa, and is abbreviated UCYN-A. The alga is the first eukaryote known to have the ability to fix nitrogen.

Reparations for slavery in the United States

From Wikipedia, the free encyclopedia

https://en.wikipedia.org/wiki/Reparations_for_slavery_in_the_United_States 
 

Reparations for slavery is the application of the concept of reparations to victims of slavery or their descendants. There are concepts for reparations in legal philosophy and reparations in transitional justice. In the US, reparations for slavery have been both given by legal ruling in court and/or given voluntarily (without court rulings) by individuals and institutions.

The first recorded case of reparations for slavery in the United States was to former slave Belinda Royall in 1783, in the form of a pension, and since then reparations continue to be proposed. To the present day, no federal reparations bills have been passed. The 1865 Special Field Orders No. 15 ("Forty acres and a mule") is the most well known attempt to help newly freed slaves integrate into society and accumulate wealth. However, President Andrew Johnson reversed this order, giving the land back to its former Confederate owners.

Reparations have been a recurring idea in the politics of the United States, most recently in the 2020 Democratic Party presidential primaries. The call for reparations intensified in 2020, amidst the protests against police brutality and the COVID-19 pandemic, which both kill Black Americans disproportionately. Calls for reparations for racism and discrimination in the US are often made by black communities and authors alongside calls for reparations for slavery. The idea of reparations remains highly controversial, due to questions of how they would be given, how much would be given, who would pay them, and who would receive them.

Forms of reparations which have been proposed in the United States by city, county, state, and national governments or private institutions include: individual monetary payments, settlements, scholarships, waiving of fees, and systemic initiatives to offset injustices, land-based compensation related to independence, apologies and acknowledgements of the injustices, token measures (such as naming a building after someone), and the removal of monuments and streets named to slave owners and defenders of slavery.

Since further injustices and discrimination have continued since slavery was outlawed in the US, some black communities and civil rights organizations have called for reparations for those injustices as well as for reparations directly related to slavery. Some suggest that the U.S. prison system, starting with the convict lease system and continuing through the present-day government-owned corporation Federal Prison Industries (UNICOR), is a modern form of legal slavery that still primarily and disproportionately affects black populations and other minorities via the war on drugs and what has been criticized as a school-to-prison pipeline.

U.S. historical context

In colonial times

The debate on reparations reaches as far back as the eighteenth century. Quakers, who were some of the first abolitionists in the United States, almost unanimously insisted that freed slaves were entitled to compensation from their former owners. If an owner repented of his sin of owning a chattel slave, he needs to atone for it by making amends. Quakers cited the book of Deuteronomy, in which owners were exhorted to share their goods with former slaves.

During the Revolutionary War, Warner Mifflin advocated for restitution for freed ex-slaves as early as 1778, in the form of cash payments, land, and shared crop arrangements. Gary B. Nash writes that, "he may fairly be called the father of American reparation".

Before the Civil War

Well before slavery was abolished nationally in 1865, abolitionists presented suggestions on what could or should be done to compensate the enslaved workers after their liberation.

Early in 1859, in a book dedicated to "Old Hero" John Brown, James Redpath declared himself a "reparation advocate", and implies that in his view, the lands of the Confederacy should be given to the ex-slaves. He also quotes an earlier poem, by William North, that refers to "the course of reparation".

Later that year, after Brown's execution, Redpath reported in the first biography of Brown that he was not merely an emancipationist, but a reparation advocate. He believed, not only that the crime of slavery should be abolished, but that reparation should be made for the wrongs that had been done to the slave. What he believed, he practiced. On this occasion [Missouri raid, 1859], after telling the slaves that they were free, he asked them how much their services had been worth, and—having been answered—proceeded to take property to the amount thus due to the negroes."

Calls for permanent confiscation and redistribution of plantation lands had already been made by Representatives George W. Julian and Thaddeus Stevens, both of the Radical Republican faction.

The Reconstruction period

The arguments surrounding reparations are based on the formal discussion about many different reparations, and actual land reparations received by African Americans which were later taken away. In 1865, after the Confederate States of America were defeated in the American Civil War, General William Tecumseh Sherman issued Special Field Orders, No. 15 to both "assure the harmony of action in the area of operations" and to solve problems caused by the masses of freed slaves, a temporary plan granting each freed family forty acres of tillable land in the sea islands and around Charleston, South Carolina for the exclusive use of black people who had been enslaved. The army also had a number of unneeded mules which were given to freed slaves. Around 40,000 freed slaves were settled on 400,000 acres (1,600 km²) in Georgia and South Carolina. However after Lincoln was assassinated, President Andrew Johnson reversed the order. The land was returned to its previous owners, and black people were forced to leave. In 1867, Thaddeus Stevens sponsored a bill for the redistribution of land to African Americans, but it did not pass.

Reconstruction came to an end in 1877 without the issue of reparations having been addressed. Thereafter, a deliberate movement of segregation and oppression arose in Southern states. Jim Crow laws were passed in some Southeastern states to reinforce the existing inequality that slavery had produced. In addition, white extremist organizations such as the Ku Klux Klan engaged in a massive campaign of terrorism throughout the Southeast in order to keep African Americans in their prescribed social place. For decades this assumed inequality and injustice was ruled on in court decisions and debated in public discourse.

In one anomalous case, a former slave named Henrietta Wood successfully sued for compensation after having been kidnapped from the free state of Ohio and sold into slavery in Mississippi. After the American Civil War, she was freed and returned to Cincinnati, where she won her case in federal court in 1878, receiving $2,500 (~$81,457 in 2024) in damages. Though the verdict was a national news story, it did not prompt any trend toward additional similar cases.

Post-Reconstruction Era

In 1896, the National Ex-Slave Mutual Relief, Bounty and Pension Association (MRB&PA) was founded for the purpose of obtaining pensions for former slaves from the Federal government as compensation and reparations for their unpaid labor and suffering. Chartered in 1898 in Nashville, Tennessee, the organization was founded by former slaves Callie House and Isaiah H. Dickerson. According to some historians, the organization was "the first mass reparations movement led by African Americans." The organization and its leaders were hounded with false allegations and criminal prosecutions until its last local branches closed in the 1930's.

In 1915, under Callie House's leadership, the association filed a class-action lawsuit, Johnson v. McAdoo, in federal court against the U.S. Treasury Department for 68 million dollars. $68 million was the amount of cotton tax collected between 1862 and 1868 and, it was argued, was due to the plaintiffs because this cotton had been produced by them and their ancestors as a result of their involuntary servitude. This was the first documented Black reparations litigation in the US on the federal level. The U.S. Court of Appeals for the District of Columbia denied the claim based on governmental immunity as did the U.S. Supreme Court, siding with the Appeals Court's decision.

2020

The topic became a prominent theme during the 2020 Democratic Party presidential primaries as concerns surrounding race were heightened due to current events. It was further amplified because of African-American people were dying prematurely and disproportionately due to the COVID-19 pandemic. Ongoing systemic racism and police brutality also sparked outrage across the country, notably the killing of Breonna Taylor, a 26-year-old African-American emergency medical technician, fatally shot by Louisville Metro Police Department in her home; the murder of Ahmaud Arbery, shot while out for a run by three white men in Georgia; and the murder of George Floyd, a Black American killed during an arrest by Minneapolis police after allegedly passing a counterfeit $20 bill, that sparked the nationwide George Floyd protests.

Candidates that endorsed the idea included:

• Andrew Yang said that he supports H.R. 40, the Commission to Study and Develop Reparation Proposals for African-Americans Act, sponsored by Rep. Sheila Jackson Lee, while speaking on the Karen Hunter show.
• Marianne Williamson detailed a plan for reparations in an interview for Ebony Magazine.
• Senators Elizabeth Warren and Cory Booker have both indicated some level of support for reparations, according to NPR.
• Tulsi Gabbard is a cosponsor of H.R. 40, the only piece of legislation in Congress to study and develop reparations proposals and Bernie Sanders is a co-sponsor for the Senate version of the bill.

Kamala Harris declared in April 2019 she supports reparations.

Beto O'Rourke is "open to considering some form of reparations," according to U.S. News & World Report.

Tom Steyer in the 2020 Democratic Primaries Debate in South Carolina voiced his support for reparations.

Proposals for reparations

United States government

Some proposals have called for direct payments from the U.S. government. Various estimates have been given if such payments were to be made. Harper's Magazine estimated that the total of reparations due was about "$97 trillion, based on 222,505,049 hours of forced labor between 1619 and 1865, regardless the United States wasn't a recognized independent country until after the Revolutionary War in 1787, compounded at 6% interest through 1993". Should all or part of this amount be paid to the descendants of slaves in the United States, the current U.S. government would only pay a fraction of that cost, since it has been in existence only since 1789. For two centuries, from the 1700s until World War I, the average wage for one day's unskilled labor in America was one dollar.

According to the Brookings Institution, in 1860, over $3 billion (~$83 billion in 2023) was the value assigned to the physical bodies of enslaved Black Americans to be used as free labor and production. This was more money than was invested in factories and railroads combined. In 1861, the value placed on cotton produced by enslaved Blacks was $250 million (~$6.66 billion in 2023). For the descendants of the 12.5 million Blacks who were shipped in chains from Western Africa, “America has a genetic birth defect when it comes to the question of race,” as stated recently by U.S. representative Hakeem Jeffries. If America is to atone for this defect, reparations for Black Americans is part of the healing and reconciliation process.

The Rev. M.J. Divine, better known as Father Divine, was one of the earliest leaders to argue clearly for "retroactive compensation", and the message was spread via International Peace Mission publications. On July 28, 1951, Father Divine issued a "peace stamp" bearing the text: "Peace! All nations and peoples who have suppressed and oppressed the under-privileged, they will be obliged to pay the African slaves and their descendants for all uncompensated servitude and for all unjust compensation, whereby they have been unjustly deprived of compensation on the account of previous condition of servitude and the present condition of servitude. This is to be accomplished in the defense of all other under-privileged subjects and must be paid retroactive up-to-date".

At the first National Reparations Convention in Chicago in 2001, a proposal by Howshua Amariel, a Chicago social activist, would require the federal government to make reparations to proven descendants of slaves. In addition, Amariel stated "For those blacks who wish to remain in America, they should receive reparations in the form of free education, free medical, free legal and free financial aid for fifty years with no taxes levied," and "For those desiring to leave America, every black person would receive a million dollars or more, backed by gold, in reparation." At the convention Amariel's proposal received approval from the 100 or so participants. Nevertheless, the question of who would receive such payments, who should pay them and in what amount, has remained highly controversial, since the United States Census does not track descent from slaves or slave owners and relies on self-reported racial categories.

On July 30, 2008, the United States House of Representatives passed a resolution apologizing for American slavery and subsequent discriminatory laws.

Nine states have officially apologized for their involvement in the enslavement of Africans. Those states are:

• Alabama – April 25, 2007
• Connecticut
• Delaware – February 11, 2016
• Florida – 2008
• Maryland – 2007
• New Jersey – 2008
• North Carolina – 2007
• Tennessee
• Virginia – 2007

Private institutions

Private institutions and corporations were also involved in slavery. On March 8, 2000, Reuters News Service reported that Deadria Farmer-Paellmann, a law school graduate, initiated a one-woman campaign making a historic demand for restitution and apologies from modern companies that played a direct role in enslaving Africans. Aetna Inc. was her first target because of their practice of writing life insurance policies on the lives of enslaved Africans with slave owners as the beneficiaries. In response to Farmer-Paellmann's demand, Aetna Inc. issued a public apology, and the "corporate restitution movement" was born.

By 2002, nine lawsuits were filed around the country coordinated by Farmer-Paellmann and the Restitution Study Group—a New York non-profit. The litigation included 20 plaintiffs, demanding restitution from 20 companies from the banking, insurance, textile, railroad, and tobacco industries. The cases were consolidated under 28 U.S.C. 1407 to multidistrict litigation in the United States District Court for the Northern District of Illinois. The district court dismissed the lawsuits with prejudice, and the claimants appealed to the United States Court of Appeals for the Seventh Circuit.

On 13th December, 2006, that court, in an opinion written by Judge Richard Posner, modified the district court's judgment to be a dismissal without prejudice, affirmed the majority of the district court's judgment, and reversed the portion of the district court's judgment dismissing the plaintiffs' consumer protection claims, remanding the case for further proceedings consistent with its opinion. Thus, the plaintiffs may bring the lawsuit again, but must clear considerable procedural and substantive hurdles first:

If one or more of the defendants violated a state law by transporting slaves in 1850, and the plaintiffs can establish standing to sue, prove the violation despite its antiquity, establish that the law was intended to provide a remedy (either directly or by providing the basis for a common law action for conspiracy, conversion, or restitution) to lawfully enslaved persons or their descendants, identify their ancestors, quantify damages incurred, and persuade the court to toll the statute of limitations, there would be no further obstacle to the grant of relief.

In October 2000, California passed the Slavery Era Disclosure Law requiring insurance companies doing business there to report on their role in slavery. The disclosure legislation, introduced by Senator Tom Hayden, is the prototype for similar laws passed in 12 states around the United States.

The NAACP has called for more of such legislation at local and corporate levels. It quotes Dennis C. Hayes, CEO of the NAACP, as saying, "Absolutely, we will be pursuing reparations from companies that have historical ties to slavery and engaging all parties to come to the table." Brown University, whose namesake family was involved in the slave trade, has also established a committee to explore the issue of reparations. In February 2007, Brown University announced a set of responses to its Steering Committee on Slavery and Justice. While in 1995 the Southern Baptist Convention apologized for the "sins" of racism, including slavery.

In December 2005, a boycott was called by a coalition of reparations groups under the sponsorship of the Restitution Study Group. The boycott targets the student loan products of banks deemed complicit in slavery—particularly those identified in the Farmer-Paellmann litigation. As part of the boycott, students are asked to choose from other banks to finance their student loans.^[65]

Pro-reparations groups such as the National Coalition of Blacks for Reparations in America advocate for compensation to be in the form of community rehabilitation and not payments to individual descendants.

Black Lives Matter

Many groups under the Black Lives Matter organization have laid out a list of demands, some of which include: reparations, for what they say are past and continuing harms to African Americans, an end to the death penalty, legislation to acknowledge the effects of slavery, a move to defund the police, seizing homes owned by white families and providing them free to blacks, as well as investments in education initiatives, mental health services, and jobs programs. These calls for reparations have been bolstered amidst the COVID-19 pandemic and the high rates of police brutality against Blacks.

Arguments for reparations

Accumulated wealth

Housing discrimination played a major role in creating the racial wealth gap that exists today. After the Great Migration of Southern blacks to Chicago in the 1940s, redlining was used to keep former slaves segregated from whites and to prevent black families from getting a mortgage. Thus they were forced to buy houses on contracts from real estate speculators, which were a scam. Not only did this cause thousands of Black Americans to lose their homes and their money, it also created what are known today as ghettos and prevented Blacks from accumulating wealth. Today, the average white family has roughly 10 times the amount of wealth as the average black family, and white college graduates have over seven times more wealth than Black college graduates.

The wealth of the United States was greatly enhanced by the exploitation of African-American slave labor: some argue it is the bedrock for the U.S. economy and capitalism. However, former slaves and their descendants are among the poorest demographic in America. According to this view, reparations would be valuable primarily as a way of correcting modern economic imbalances.

In 2008 the American Humanist Association published an article which argued that if emancipated slaves had been allowed to possess and retain the profits of their labor, their descendants might now control a much larger share of American social and monetary wealth. Not only did the freedmen not receive a share of these profits, but they were stripped of the small amounts of compensation paid to some of them during Reconstruction.^[71] Therefore, many scholars and activists call for reparations to eliminate "racial disparities in wealth, income, education, health, sentencing and incarceration, political participation, and subsequent opportunities to engage in American political and social life".

Health care

In 2019, VICE magazine published an article that argued racial health disparities, from slavery through Jim Crow until today, have cost Black Americans a significant amount of money in health care expenses and lost wages, and should be paid back. Ray and Perry state in a Brookings article that the lack of a social safety net and the wealth gap are particularly highlighted during the COVID-19 pandemic. They explain that “disparities in access to health care along with inequities in economic policies combine,” making this inequality a life or death situation for black Americans.

Current discrimination

Many argue that giving reparations for slavery is too complicated, but there is a strong basis for them on the past and current discrimination that blacks in America face. Ta-Nehisi Coates explains it in "The Case for Reparations" article in The Atlantic as "ninety years of Jim Crow, sixty years of separate but equal, and thirty-five years of racist housing policy". The legacy of these policies have kept African Americans from opportunities to build wealth, while slavery "enriched white slave owners and their descendants". Today, the district of North Lawndale in Chicago, where redlining was the strongest, is the poorest neighborhood in the city, with an unemployment rate of 18.6% and 42% of residents living below the poverty line.

The discriminatory practices of 1940 through 1970 still reverberate today, as the average White family has roughly ten times the amount of wealth as the average Black family. As Bittker claims in his book The Case for Black Reparations, "as slavery faded into the background, it was succeeded by a caste system embodying white supremacy". Many argue that while reparations may be a first step towards amending the harms caused by slavery, the systemic racism that exists in many institutions will not be fixed as easily. Malcolm X stated: "If you stick a knife in my back nine inches and pull it out six inches, there's no progress. If you pull it all the way out that's not progress. Progress is healing the wound that the blow made."

Precedents

Advocates have used other examples of reparations to argue that victims of institutional slavery should be similarly compensated.

In several cases the federal government has formally apologized to or compensated minority groups for past actions:

• Under the Civil Liberties Act of 1988, signed into law by President Ronald Reagan, the U.S. government apologized for Japanese American internment during World War II and provided reparations of $20,000 to each survivor, to compensate for loss of property and liberty during that period. No compensation was given to the descendants of affected individuals though.
• The Alaska Native Claims Settlement Act transferred land, federal money, and a portion of oil revenues to native Alaskans.
• The Apology Resolution of 1993 apologized for the overthrow of the Kingdom of Hawaii, but gave no compensation.

U.S. state governments have made reparations in some specific circumstances:

• Virginia established a compensation fund for victims of involuntary sterilization in 2015.

Other countries have also opted to pay reparations for past grievances, such as:

• Reparations for the Holocaust, including the Reparations Agreement between Israel and West Germany and various programs under the Conference on Jewish Material Claims Against Germany.

Arguments against reparations

Statute of limitations

Most state and federal laws under which parties can sue for damages have a statute of limitations which sets a deadline for filing; these have all long since passed, which prevents courts from granting relief under existing laws. This has been used effectively in several suits, including In re: African American Slave Descendants, which dismissed a high-profile suit against a number of businesses with ties to slavery.

Technical complications

The technical side of reparations is very complex, and could be a reason why they have not yet been implemented. Some argue against the idea of putting a monetary value on the traumas that Black Americans have faced, dubbing it "transactionalism". On the other hand, some dismiss the case for reparations entirely due to practical concerns, such as who would receive these financial payments, why should the current generation pay for wrongs for which they are not responsible, and how much should be paid.

The estimates of the monetary value of stolen slave labor and subsequent discrimination vary “from an outrageously low $3.2 million to $4.7 billion,” and to as much as $12 trillion. This also raises the question of who is responsible for paying. Generally, three actors are agreed upon: federal and state governments, who supported and protected the institution of slavery; private companies that benefited from it; and “rich families that owe a good portion of their wealth to slavery”.

Some claim that closing the wealth gap involves paying descendants of slaves “individual cash payments in the amount that will close the Black-white racial wealth divide”. Another suggestion is for reparations to "come in the form of wealth-building opportunities that address racial disparities in education, housing, and business ownership". For example, in the city of Asheville, North Carolina, reparations have been implemented in the form of "investments in areas where Black residents face disparities". However, the complications that surround this are significant, and others argue that putting the money into communities is not efficient, due to people moving and gentrification.

In his book, Bittker lays out some of the practical and constitutional problems that would likely arise in an attempt to execute a program of reparations to Blacks. Would it be the same payment to every person? Would they have to prove ancestry to an African slave, or would it be any black person who was subject to racism? There are no real answers to these questions, as this is an unprecedented case. Other cases of reparations, such as to the Jewish people who survived the Holocaust or the Native Americans in the United States, are very different in the way that it is much easier to identify the group who should receive them, and the reparations were paid more quickly than in the case of reparations for slavery.

Additional arguments and opinions

Steven Greenhut, the western region director for the R Street Institute, has suggested that reparations would make racism worse.

Republican senator Mitch McConnell of Kentucky, who is a descendant of slave owners, while acknowledging that slavery was an "original sin" of the United States, opposes providing reparations because he believes "none of us currently living are responsible."

One publication against reparations is David Horowitz, Uncivil Wars: The Controversy Over Reparations for Slavery (2002). Other works that discuss problems with reparations include John Torpey's Making Whole What Has Been Smashed: On Reparations Politics (2006), Alfred Brophy's Reparations Pro and Con (2006), and Nahshon Perez's Freedom from Past Injustices (Edinburgh University Press, 2012).

Reparations in the U.S. have never gained widespread public support. Often in these conversations, the White reaction is to claim that this is a form of unjustifiable "reverse racism", or that demands for reparations are an example of the "Black refusal to move beyond the memory of slavery". A 2020 poll from The Washington Post showed that "63% of Americans don't think the U.S. should pay reparations to the descendants of slaves". Notably, 82% of Black Americans support reparations, while 75% of White Americans do not. Some arguments also highlight the complications behind reparations, such as "not all Black Americans are descendants of slaves" or that the people alive today are not responsible for the harms of slavery. Others still argue that reparations will do nothing in the face of racism, and that structural and policy changes would be more effective. In the midst of America's racial unrest from 2020 to the present, these tensions were particularly exposed.

Reparations and COVID-19

The call for reparations has amplified due to the coronavirus pandemic, with people of color disproportionately likely to be laid off, to struggle financially, and to die from the virus. For example, 40% of black-owned businesses have closed permanently since March due to the pandemic, compared to 17% of white-owned businesses during the same period. This relates back to the fact that white families have roughly ten times the wealth of black families. This limits black-owned businesses' access to credit and loans, and they do not have the safety net in times of crises that many white-owned businesses do.

In addition, African Americans continue to get infected and die from COVID-19 at rates more than 1.5 times their share of the population. In August 2020, the CDC released data showing that Blacks, Latinos, and American Indians are experiencing hospitalizations at rates 4.5 to 5.5 times higher than non-Hispanic whites, and that African Americans are dying at 2.4 times the white rate.

Legislation and other actions

Federal government

On July 30, 2008, the United States House of Representatives passed a resolution apologizing for American slavery and subsequent discriminatory laws. The Senate apologized in 2009.

States

Legislation

• California – Adopted legislation requiring insurance companies to determine whether they have records going back to when slavery existed in this country and, if so, to provide information on insurance policies held by slaveholders on slaves to the state's insurance department. The California Reparations Task Force was established in 2020 as a non-regulatory state agency to study and develop reparation proposals.
• Illinois – Adopted legislation requiring insurance companies to determine whether they have records going back to when slavery existed in this country and, if so, to provide information on insurance policies held by slaveholders on slaves to the state's insurance department.
• Iowa – Adopted legislation asking the insurance commissioner to request if insurance companies they have records going back to when slavery existed in this country and, if so, to provide information on insurance policies held by slaveholders on slaves to the state's insurance department.
• Maryland – Adopted legislation requiring insurance companies to determine whether they have records going back to when slavery existed in this country and, if so, to provide information on insurance policies held by slaveholders on slaves to the state's insurance department.
• New York – In December of 2023, Gov. Kathy Hochul signed a bill setting up a commission for the study of slavery's legacy. The commission will possibly make recommendations for monetary compensation or other reparations for New York's Black residents under the effort. The Democrat-controlled state Legislature approved the measure in June.

Apologies

• Alabama – Apologized for its involvement in the enslavement of Africans on April 25, 2007.
• Connecticut – In 2009 apologized for its involvement in the enslavement of Africans.
• Delaware – Apologized for its involvement in the enslavement of Africans on February 11, 2016.
• Florida – In 2008, apologized for its involvement in the enslavement of Africans in America.
• Maryland – In 2007, apologized for its involvement in the enslavement of Africans in America.
• New Jersey – In 2007, apologized for its involvement in the enslavement of Africans in America.
• North Carolina – In 2007, apologized for its involvement in the enslavement of Africans in America.
• Tennessee – In 2007, the Tennessee House of Representatives voted in unanimous support on a resolution stating that it "regrets" its involvement in the enslavement of Africans. The House had specifically removed any "apology" language from the resolution.
• Virginia – Apologized for its involvement in the enslavement of Africans on February 26, 2007.

Counties

• Buncombe County, North Carolina: On June 16, 2020, in a 7–0 vote, Buncombe County Commissioners decided to remove several Confederate monuments including the Vance Monument which is named after North Carolina governor Zeb Vance, a slave owner who used convict labor to build the railroad to Western North Carolina. Significant community involvement led to the decision. Leading up to the vote, the board received 549 supporting messages and 19 opposing.

Cities

• Chicago, Illinois: "In 2015, Chicago enacted a reparations ordinance covering hundreds of African Americans tortured by police from the 1970s to the 1990s. The law calls for $5.5 million in financial compensation, as well as hundreds of thousands more for a public memorial, and a range of assistance related to health, education and emotional well-being."
• Evanston, Illinois: "The City Council of Evanston, Illinois, voted to allocate the first $10 million in tax revenue from the sale of recreational marijuana (which became legal in the state on January 1, 2020) to fund reparations initiatives that address the gaps in wealth and opportunity of black residents."
• Asheville, North Carolina: The city council approved reparations on a 7–0 vote on July 14, 2020. "[B]udgetary and programmatic priorities may include but not be limited to increasing minority home ownership and access to other affordable housing, increasing minority business ownership and career opportunities, strategies to grow equity and generational wealth, closing the gaps in health care, education, employment and pay, neighborhood safety and fairness within criminal justice," the resolution reads. The resolution establishes the Community Reparations Commission which will make concrete recommendations for programs and resources allocations to ultimately carry out the reparations. The Asheville City Council also voted unanimously on June 9, 2020, to remove two confederate monuments as a result of demands made by a group called "Black Asheville Demands" and the work of the Racial Justice Coalition with led the push for the effort. The city council meeting had so much community engagement public comment was extended for an extra hour beyond the normal meeting time.
• San Francisco, California: In March 2023, "reparation payments of $5 million to eligible Black residents [were] unanimously accepted by San Francisco’s Board of Supervisors."

Organizations and institutions

Aetna: Apologized in 2000 for issuing life insurance policies to enslavers covering the lives of enslaved people from c. 1853 to 1860. In 2002, when Farmer-Paellman brought suit against Aetna, CSX and Fleet for unjust enrichment by "a system that enslaved, tortured, starved and exploited human beings," this suit was dismissed.

University of Alabama: Apologized for the history of slavery at the university in 2004.

Georgetown University: "In 2016 [the university agreed] to give admissions preference to descendants of the 272 slaves[,] formally apologized for its role in slavery [and] [renamed] two buildings on its campus to acknowledge the lives of enslaved people". In April, 2019 students at Georgetown University voted to increase their tuition by $27.20 to benefit the descendants of the 272 slaves sold by the Jesuits who ran the school in 1838. The student-led referendum was non-binding. Later that year, after further pressure and follow up from the Georgetown University Student Association, the university eventually moved forward with a similar proposal without the students' covering the cost with a tuition increase.

Harvard University started to research its relationship to slavery in 2007. In 2016 Harvard president Drew Faust acknowledged that the university had been "directly complicit" in slavery. In 2022 the university published Harvard & the Legacy of Slavery, more than a hundred pages long. The report included recommendations to develop educational partnerships with historically Black colleges and universities, create a public memorial, and identify living descendants of people enslaved by university staff, leaders and faculty. There was a US$100m endowment for implementation. The university has given descendants of slaves preferential consideration for admissions and set up a reconciliation fund for them.

JP Morgan Chase: Apologized for its connection to slavery in 2005.

Princeton Theological Seminary: In 2019 the Seminary announced a $27 million commitment for various initiatives to recognize how it benefited from black slavery. This is the largest monetary commitment by an educational institution.

Virginia Theological Seminary: Set aside $1.7 million to pay reparations to descendants of African Americans who were enslaved to work on their campus, first distributed in 2021.

Wachovia: Apologized for its connection to slavery in 2005.