When we first met Neanderthals

Author : Wahid Ahmad

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What makes us truly human? Is it our upright, hairless bodies or our highly organized brains, our sophisticated culture, or our ability to articulate complex ideas through precise symbols? Fascinatingly, we share many of these remarkable traits with Neanderthals, our closest and most intriguing evolutionary cousins. Yet, many anthropologists argue that Neanderthals are a distinct species, pointing to unique physical characteristics. While modern humans have a rounded braincase, a small divided brow ridge, and a narrow pelvis, Neanderthals display a longer skull, a prominent continuous brow ridge, and wider hipbones. These differences are significant to categorise Neanderthals and modern humans as separate species. Genetic data indicates that our lineages diverged over 500,000 years ago, though we occasionally interbred, leaving a fascinating legacy in our shared history.

Until a few years ago, scientists thought that while Homo sapiens started spreading out from Africa at least 60,000 years ago, they didn't reach Europe until much later, around 41,000 years ago. This delay was believed to be due to the need for adaptations to Europe's colder climate or because Neanderthals successfully kept modern humans out for many millennia.

However, recent research has shown that early Homo sapiens and late Neanderthals likely coexisted in Europe for a longer period than previously thought. Additionally, DNA evidence reveals that these two groups interbred multiple times both in Europe and Asia.

Modern human genetic diversity patterns have been shaped by past demographic events and interactions. As humans migrated out of Africa, a series of events like geographic isolations, population replacement, gene flow and selective pressures created a gradient of genetic diversity, with populations farther from Africa genetically less diverse.  

Recent research highlights that modern humans also carry genetic legacies from admixture with archaic humans, including Neanderthals and Denisovans. Neanderthal ancestry is found in Eurasia, Oceania, the Americas, and North Africa, while Denisovan ancestry is mainly in Asia, the Americas, and Oceania. Additional archaic ancestry has been identified in African populations. Archaic ancestry levels vary with about 1% in Africans, 2% in Eurasians, and up to 6% in Oceania.

Neanderthals evolved in Europe for at least 400,000 years. In the Atapuerca hills of northern Spain, the Sima de los Huesos cave contains many early human skeletons dating from about 430,000 years ago. These bones were identified as early Neanderthals, a conclusion supported by ancient DNA analysis.

Genetic data from these fossils and later Neanderthals suggest that Neanderthals and modern humans began to diverge about 600,000 years ago. This challenges previous beliefs about the last common ancestor of Neanderthals and Homo sapiens, which was thought to be Homo heidelbergensis or rhodesiensis. Fossils of this species are found in Europe and Africa and were believed to have started splitting into Neanderthals and modern humans about 500,000 years ago.

However, new dating of the Kabwe cranium belonging to Homo rhodesiensis indicates it is only about 300,000 years old, much younger than expected. Additionally, studies show that rhodesiensis had a facial structure less likely to be ancestral to modern humans. Therefore, there isn't enough evidence to determine the exact nature or location of our last common ancestor with Neanderthals from around 600,000 years ago.

If our evolution goes back about 600,000 years, where are the early fossils of our species? Until recently, scientists thought the oldest Homo sapiens fossils were from Ethiopia, dating back 150,000 to 200,000 years. These fossils named Omo Kibish  and Herto, have features typical of modern humans. However, there is a large time gap between these fossils and our ancient common ancestor with Neanderthals.

In 2017, fossils from Jebel Irhoud in Morocco, dating back 300,000 years, suggested early members of the Homo sapiens lineage. These fossils have a mix of ancestral and more modern features, along with evidence of complex behaviors like controlled use of fire and advanced stone tools.

At around 300,000 years ago, there were other human species in Africa like Homo naledi in South Africa.

Pan-African model of human origin is now widely accepted, as early Homo sapiens were diverse and widely spread across Africa. Changing climates caused different populations to evolve, merge, or disappear over hundreds of thousands of years, eventually leading to modern humans.

About 60,000 years ago, Homo sapiens began to spread from Africa, with Neanderthals disappearing in the following 20,000 years of the dispersal. New discoveries show that these two events may be connected, revealing interactions and interbreeding between the species 40,000 to 60,000 years ago.

Although Neanderthals and Homo sapiens evolved mainly in separate regions, they exchanged some genes about some 250,000 years ago, possibly due to brief migrations of early Homo sapiens into Eurasia. Evidence of this contact includes shared technology and fossils like those found in Apidima Cave from Greece. One sapiens-like braincase from this site is dated to at least 210,000 years ago.

This fossil called Apidima 1 has features typical of Homo sapiens, such as a high and rounded braincase, suggesting an early population of Homo sapiens in Greece by 210,000 years ago. This group was likely related to similar populations in the Levant and was later replaced by Neanderthals around 170,000 years ago.

If the Apidima analyses are correct, Homo sapiens entered Europe over 150,000 years earlier than previously thought. The most likely route from Africa was through the Levant and Turkey. Evidence of early DNA exchanges between Neanderthals and Homo sapiens supports this theory.

There are signs of other early excursions of Homo sapiens from Africa, such as fossils in Israel at sites like Skhul, Qafzeh, and Misliya dating over 100,000 years ago. These remains show that later Homo sapiens were present in the region between 100,000 to 130,000 years ago.

Recent finds at Nesher Ramla in Israel suggest there was significant human variation in the region, possibly including co-existence of Homo sapiens and Homo neanderthalensis.

Interbreeding with archaic humans introduced new genetic variants into modern humans. Over time selective forces shaped these archaic genome segments, influencing their current distribution among modern humans.

Neanderthal DNA makes up about 2% of the genomes of all non-African populations today. Different individuals carry different Neanderthal variants, and these variants can vary significantly in frequency among populations. A smaller number of Neanderthal-derived variants are also found in sub-Saharan Africa due to gene flow from Europe and Western Asia after the Neanderthals disappeared.

The consistent 2% Neanderthal DNA in present-day genomes suggests that most Neanderthal contributions occurred about 60,000 years ago when populations left Africa and became ancestors to all non-Africans. However, some Neanderthals also contributed locally to early European modern human groups, such as a 40,000-year-old individual from Romania and three 45,000-year-old individuals from Bulgaria. These early populations did not leave many descendants that contributed genetic variants to present-day populations, so their impact was limited.

The genetic changes that accumulated on archaic and modern human lineages impacted the physiology of Neanderthals, Denisovans, and modern humans, with the first two becoming extinct.

When modern humans interbred with Neanderthals, their offspring had one set of modern human chromosomes and one set of archaic chromosomes. Over 2,000 generations, these archaic DNA segments became shorter due to recombination, resulting in scattered fragments in today's genomes.

Typically, an archaic DNA segment today is about 50 kilobases long. However, the actual lengths vary, reflecting past recombination events. For a fragment to be confidently identified as archaic and from about 2,000 generations ago, it must be of substantial length.

Some DNA segments in humans are similar to Neanderthal or Denisovan DNA not because of recent gene flow, but because these segments have persisted since the common ancestor of modern and archaic humans. These segments are usually shorter than 12 kb due to longer recombination periods.

The size of archaic DNA fragments also depends on local recombination rates, which vary across the genome, among populations, and over time.

A typical 50 kb segment of Neanderthal DNA will carry about 16 unique variants from mutations that occurred on the Neanderthal lineage. Modern humans have accumulated a similar number of unique variants independently. Thus, archaic DNA segments often stand out because they carry more single nucleotide variants than surrounding regions.

The way Neanderthal segments are spread across modern human genomes suggests interbreeding occurred between 50,000 and 60,000 years ago, before East Asians and Europeans split into distinct groups.

The genome of an ancient individual called Ust’-Ishim, who is equally related to modern East Asians and Europeans, has similar levels of Neanderthal DNA as present-day Eurasians. This supports a single episode of interbreeding around 52,000–58,000 years ago.

Several ideas try to explain why East Asians have more Neanderthal DNA. It could be due to stronger natural selection in Europe reducing Neanderthal DNA over time, or because a population without Neanderthal genes mixed with Europeans later on. Another possibility is that interbreeding happened multiple times: initially into the ancestors of both East Asians and Europeans, followed by additional mixing with East Asians after the populations had diverged.

Scientists analysed Neanderthal DNA fragments in modern human genomes, found that East Asians have more Neanderthal ancestry compared to Europeans with arround 19.6% enrichment in East Asians.

By studying Neanderthal DNA fragments in Europeans and East Asians, strong evidence supporting multiple episodes of interbreeding between Neanderthals and humans.

After an initial mixture when humans first entered Eurasia, there were subsequent pulses of Neanderthal DNA introgression into both European and East Asian populations. This on-going mixing likely contributed to the higher level of Neanderthal ancestry observed in East Asians compared to Europeans.

 

These findings contrast with theories proposing that differences in Neanderthal ancestry between Europe and Asia are solely due to dilution in Europe from a population called basal Eurasians, who lacked Neanderthal ancestry. While dilution may have played a role, results indicate that multiple admixture events were crucial in shaping Neanderthal DNA patterns across Eurasia.

Neandertal genes in modern humans can affect how our bodies process food and drugs. For instance, some Neandertal DNA on chromosome 17 alters how a liver protein handles fats and sugars, making people more prone to type 2 diabetes. Other Neandertal genes increase the risk of protein and calorie deficiencies and can affect how medications are metabolized in the body.

Neandertal genes affect how we feel pain and handle pregnancies today. On chromosome 2, a piece of Neandertal DNA changes a protein (SCN9A) that controls how sensitive we are to pain. Neandertals had versions of this protein that made them more sensitive to pain. Today, about 0.4% of people in the UK carry this Neandertal version and say they feel more pain than others. This sensitivity might have helped Neandertals avoid danger.

On chromosome 11, another piece of Neandertal DNA affects the progesterone receptor, important for pregnancy. This gene variant is linked to a higher risk of premature in modern humans. But it also seems to protect against bleeding and miscarriages early in pregnancy. This trade-off suggests that these genetic differences from Neandertals have influenced our health today, showing how ancient genes still play a role in our lives. Notably, two different versions of the Neandertal progesterone receptor gene have been contributed to modern humans, and both have risen in frequency, as shown by an increase in their occurrence in skeletal remains of individuals over the past 10,000 years. Both Neandertal versions result in a higher progesterone effect during pregnancies. This is compatible with the finding that progesterone administration lowers miscarriage rates in women who previously experienced miscarriages.

Archaic humans like Neandertals left genetic marks in our immune systems that still affect us today. They had gene variants that helped them fight infections, likely because diseases were a big threat back then. Some of these Neandertal genes are still with us and influence how our immune systems respond to infections.

For example, on chromosome 4, there's a DNA segment from Neandertals that boosts our immune response by increasing the expression of receptors on cells that recognize and fight microbes. This likely helped Neandertals resist certain infections, like Helicobacter pylori. Today, these genetic variants are found in different frequencies across human populations, which suggests they were influenced by local diseases over time.

Another Neandertal DNA segment on chromosome 3 affects our risk from severe infections like SARS-CoV-2. People with this segment have a higher risk of needing mechanical ventilation or dying from such infections. However, this segment also reduces the risk of getting infected with HIV, possibly by affecting the expression of a gene called CCR5.

These genetic differences show how ancient humans adapted to their environments, including the diseases they faced. It's similar to how modern humans adapted to diseases like malaria, which shaped the prevalence of traits like sickle cell anemia. Studying these Neandertal genes gives us insights into our evolutionary history and why our immune systems vary across different populations today.

Complex traits in humans, like height or cognitive abilities, are influenced by many genetic variants scattered across the genome, as well as environmental factors. Understanding how genes from archaic humans affect such traits is challenging because each genetic change typically has a small effect.

Researchers don't directly link individual archaic variants to specific traits like height or intelligence. Instead, they analyze whether the collective presence of Neandertal or Denisovan genes in a population tends to influence traits in a certain direction or contributes to the variability in the risk of complex diseases.

For instance, studies have found that Neandertal DNA contributes significantly to variations in the risk of conditions like depression and sun-induced skin lesions. However, when looking across a wide range of traits, it's been observed that archaic genes have less impact than initially expected. Among 405 complex traits studied, dermatological traits showed the most influence from Neandertal ancestry, while cognitive traits were least affected. This aligns with the understanding that Neandertal genetic influence is less pronounced in brain-related genes compared to genes influencing other parts of the body.

Gene expression levels play a crucial role in influencing many complex traits. Studies have shown that Neandertal genetic variants tend to be expressed at lower levels compared to modern human variants in individuals who carry both types of genes. This difference is particularly noticeable in tissues like the testis and parts of the brain such as the cerebellum and basal ganglia. It suggests that there has been strong evolutionary pressure against Neandertal regulatory sequences in these specific tissues, which aligns with the limited impact of Neandertal genes on cognitive traits.

 

Archaic variants that are older seem to have been more tolerated among modern humans compared to more recent ones. Variants shared between different Neandertal individuals or between Neandertals and Denisovans are likely older. In terms of gene regulation, older archaic variants that increase gene expression are less common in present-day populations, except for those shared between Neandertals and Denisovans. This suggests that variants persisting in archaic groups for longer periods might have had less harmful effects on modern humans, possibly because they were better adapted to ancestral conditions or underwent more rigorous selection processes over time.

Denisovans are less studied due to the limited availability of high-quality genomes. However, one significant Denisovan genetic contribution is a 33-kb DNA segment on chromosome 2 found in over 80% of Tibetans but rare in other Asian populations. This segment contains the EPAS1 gene, which helps in adapting to low oxygen levels at high altitudes. Denisovans likely adapted to life on the Tibetan plateau and passed on this genetic advantage to modern humans who migrated to the region.

 

Genetic variants inherited from Neandertals affect how genes are turned on and off in modern humans. Studies show that Neandertal variants tend to be expressed less than modern human variants, especially in tissues like the brain and testis. Older Neandertal variants are better tolerated in modern populations, possibly because they had more time to undergo natural selection. Newer variants may have more detrimental effects since they haven't been refined through evolution as much. These insights help explain how Neandertal genes influence traits and diseases in present-day humans.

Ancestral genetic variants that were once thought absent in present-day humans are now being discovered in diverse populations worldwide. These variants either persisted since the common ancestors of modern and archaic humans around 600,000 years ago, especially prevalent in African populations with high genetic diversity, or were introduced by interbreeding with Neandertals and Denisovans. Populations like those in Oceania, with higher levels of archaic ancestry, show particularly elevated levels of reintroduced ancestral variants. As genomic studies expand to include more diverse populations, more of these ancient genetic variants, previously considered lost, are likely to be uncovered. This broader data will enhance our understanding of how these variants influence human traits and health.

The presence of rare ancestral genetic variants in modern humans challenges how we define modernity genetically. Paleoanthropologists define Neandertals based on skeletal features like robustness, prominent brow ridges, and certain skull shapes, most of which are distinctive to Neandertals. However, some modern humans exhibit similar skeletal traits. Yet, only Neandertals possess a combination of all or most of these features.

Similarly, modern human genetics can be understood as a mix of derived genetic features, not universally present in all individuals today. Some modern traits exist in their ancestral forms in certain people due to persistence from common ancestors with archaic humans or through interbreeding with Neandertals and Denisovans. Conversely, some modern genetic traits are found in late archaic humans, showing they were introduced from modern humans.

Certain modern human traits are nearly fixed across all populations, such as specific genetic changes that are absent in Neandertals and Denisovans. Studying these unique genomic regions can provide insights into what makes modern humans distinct genetically.

The genomes of Neandertals and Denisovans reveal they shared common ancestors with modern humans around 600,000 years ago, while the ancestor of all present-day humans existed about 300,000 years ago. Only a small fraction of the genome, about 1.5% to 7%, differs significantly between modern and archaic humans. These regions contain genetic variants where frequencies vary greatly between the two groups, influencing the modern human phenotype.

 

From a genetic standpoint, modern humans are defined by a combination of genetic features, where each person carries most but not necessarily all of these traits. This "combinatorial view" acknowledges that some genetic changes may have had more significant functional impacts than others. The emergence of these changes likely occurred in a population ancestral to modern humans, setting them apart from other contemporaneous human forms.

 

Studying gene flow between archaic and modern humans helps us understand the physiological effects of these genetic variants. Archaic variants introduced into modern human populations through interbreeding are often rare, making them challenging to study. Increasing diversity in biobanks, with participants from various ancestries, will be crucial for studying these rare variants, as they may have reached higher frequencies in smaller, isolated populations.

It's essential to recognize that ancestral variants associated with developmental or adult effects are not inherently "primitive" or "pathological." These variants functioned effectively in healthy archaic humans for hundreds of thousands of years, closely related to modern humans, and likely continue to function well in people today.

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