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One of the best ways to study human evolution is by comparing us with nonhuman species that, evolutionarily speaking, are closely related to us. That closeness can help scientists narrow down precisely what makes us human, but that scope is so narrow it can also be extremely hard to define. To address this complication, researchers from Stanford University have developed a new technique for comparing genetic differences.

An image, from previous research, of human cortical spheroids derived in the lab of Sergiu Pașca, associate professor of psychiatry and behavioral sciences. (Image credit: Timothy Archibald)

Through two separate sets of experiments with this technique, the researchers discovered new genetic differences between humans and chimpanzees. They found a significant disparity in the expression of the gene SSTR2 – which modulates the activity of neurons in the cerebral cortex and has been linked, in humans, to certain neuropsychiatric diseases such as Alzheimer’s dementia and schizophrenia – and the gene EVC2 , which is related to facial shape. The results were published March 17 in Nature and Nature Genetics , respectively.

“It’s important to study human evolution, not only to understand where we came from, but also why humans get so many diseases that aren’t seen in other species,” said Rachel Agoglia, a recent Stanford genetics graduate student who is lead author of the Nature paper.

The Nature paper details the new technique, which involves fusing human and chimpanzee skin cells that had been modified to act like stem cells – highly malleable cells that can be prodded to transform into a variety of other cell types (albeit not a full organism).

“These cells serve a very important specific purpose in this type of study by allowing us to precisely compare human and chimpanzee genes and their activities side-by-side,” said Hunter Fraser, associate professor of biology at Stanford’s  School of Humanities and Sciences . Fraser is senior author of the Nature Genetics paper and co-senior author of the Nature paper with Sergiu Pașca, associate professor of psychiatry and behavioral sciences in the Stanford  School of Medicine .

Close comparisons

The Fraser lab is particularly interested in how the genetics of humans and other primates compare at the level of cis-regulatory elements, which affect the expression of nearby genes (located on the same DNA molecule, or chromosome). The alternative – called trans-regulatory factors – can regulate the expression of distant genes on other chromosomes elsewhere in the genome. Due to their broad effects, trans-regulatory factors (such as proteins) are less likely to differ among closely related species than cis-regulatory elements.

But even when scientists have access to similar cells from humans and chimpanzees, there is a risk of confounding factors. For example, differences in the timing of development between species is a significant hurdle in studying brain development, explained Pașca. This is because human brains and chimpanzee brains develop at very different rates and there is no exact way to directly compare them. By housing human and chimpanzee DNA within the same cellular nucleus, scientists can exclude most confounding factors.

For the initial experiments using these cells, Agoglia coaxed the cells into forming so-called cortical spheroids or organoids – a bundle of brain cells that closely mimics a developing mammalian cerebral cortex. The Pașca lab has been at the forefront of developing brain organoids and assembloids for the purpose of researching how the human brain is assembled and how this process goes awry in disease.

“The human brain is essentially inaccessible at the molecular and cellular level for most of its development, so we introduced cortical spheroids to help us gain access to these important processes,” said Pașca, who is also the Bonnie Uytengsu and Family Director of Stanford Brain Organogenesis.

As the 3D clusters of brain cells develop and mature in a dish, their genetic activity mimics what happens in early neurodevelopment in each species. Because the human and chimpanzee DNA are bound together in the same cellular environment, they are exposed to the same conditions and mature in parallel. Therefore, any observed differences in the genetic activity of the two can reasonably be attributed to actual genetic differences between our two species.

Through studying brain organoids derived from the fused cells that were grown for 200 days, the researchers found thousands of genes that showed cis-regulatory differences between species. They decided to further investigate one of these genes – SSTR2 – which was more strongly expressed in human neurons and functions as a receptor for a neurotransmitter called somatostatin. In subsequent comparisons between human and chimpanzee cells, the researchers confirmed this elevated protein expression of SSTR2 in human cortical cells. Further, when the researchers exposed the chimpanzee cells and human cells to a small molecule drug that binds to SSTR2 , they found that human neurons responded much more to the drug than the chimpanzee cells.

This suggests a way by which the activity of human neurons in cortical circuits can be modified by neurotransmitters. Interestingly, this neuromodulatory activity may also be related to disease since SSTR2 has been shown to be involved in brain disease.

“Evolution of the primate brain may have involved adding sophisticated neuromodulatory features to neural circuits, which under certain conditions can be perturbed and increase susceptibility to neuropsychiatric disease,” said Pașca.

Fraser said these results are essentially “a proof of concept that the activity we’re seeing in these fused cells is actually relevant for cellular physiology.”

Investigating extreme differences

For the experiments published in Nature Genetics , the team coaxed their fused cells into cranial neural crest cells, which give rise to bones and cartilage in the skull and face, and determine facial appearance.

“We were interested in these types of cells because facial differences are considered some of the most extreme anatomical differences between humans and chimps – and these differences actually affect other aspects of our behavior and evolution, like feeding, our senses, brain expansion and speech,” said David Gokhman, a postdoctoral scholar in the Fraser lab and lead author of the Nature Genetics paper. “Also, the most common congenital diseases in humans are related to facial structure.”

In the fused cells, the researchers identified a gene expression pathway that is much more active in the chimpanzee genes of the cells than in the human genes – with one specific gene, called EVC2 , appearing to be six times more active in chimpanzees. Existing research has shown that people who have inactive EVC2 genes have flatter faces than others, suggesting that this gene could explain why humans have flatter faces than other primates.

What’s more, the researchers determined that 25 observable facial features associated with inactive EVC2 are noticeably different between humans and chimpanzees – and 23 of those are different in the direction the researchers would have predicted, given lower EVC2 activity in humans. In follow-up experiments, where the researchers reduced the activity of EVC2 in mice, the rodents, too, developed flatter faces.

Another tool in the toolbox

This new experimental platform is not intended to replace existing cell comparison studies, but the researchers hope it will support many new findings about human evolution, and evolution in general.

“Human development and the human genome have been very well studied,” said Fraser. “My lab is very interested in human evolution, but, because we can build on such a wealth of knowledge, this work can also reveal new insights into the process of evolution more broadly.”

Looking forward, the Fraser lab is working on differentiating the fused cells into other cell types, such as muscle cells, other types of neurons, skin cells and cartilage to expand their studies of uniquely human traits. The Pașca lab, meanwhile, is interested in investigating genetic dissimilarities related to astrocytes – large, multi-functional cells in the central nervous system often overlooked by scientists in favor of the flashier neurons.

“While people often think about how neurons have evolved, we should not underestimate how astrocytes have changed during evolution. The size difference alone, between human astrocytes and astrocytes in other primates, is massive,” said Pașca. “My mentor, the late Ben Barres, called astrocytes ‘the basis of humanity’ and we absolutely think he was onto something.”

Additional Stanford co-authors for the Nature paper are former research assistant Danqiong Sun, postdoctoral scholar Fikri Birey, senior research scientist Se-Jin Yoon, postdoctoral scholar Yuki Miura and former research associate Karen Sabatini.

This work was funded by a Stanford Bio-X Interdisciplinary Initiatives Seed Grant, the National Institutes of Health, the Department of Defense, the Stanford Center for Computational, Evolutionary and Human Genomics, the Stanford Medicine’s Dean’s Fellowship, MCHRI, the American Epilepsy Society, the Stanford Wu Tsai Neurosciences Institute’s Big Idea Grants on Brain Rejuvenation and Human Brain Organogenesis, the Kwan Research Fund, the New York Stem Cell Robertson Investigator Award, and the Chan Zuckerberg Ben Barres Investigator Award.

Additional Stanford co-authors for the Nature Genetics paper are graduate student Maia Kinnebrew; former undergraduate Wei Gordon; former technician Danqiong Sun; postdoctoral research fellows Vivek Bajpai and Sahin Naqvi; Dmitri Petrov, the Michelle and Kevin Douglas Professor in the School of Humanities and Sciences; Joanna Wysocka, the Lorry Lokey Professor and professor of developmental biology; and Rajat Rohatgi, associate professor of biochemistry and of medicine. Researchers from University of California, San Francisco; University of Michigan, Ann Arbor; Yerkes National Primate Research Center; Emory University School of Medicine; and University of Pennsylvania are also co-authors.

This work was funded by the Human Frontier, Rothschild and Zuckerman fellowships, and the National Institutes of Health.

Fraser is a member of  Stanford Bio-X , the Maternal & Child Health Research Institute (MCHRI) , and the Stanford Cancer Institute . Pașca is a member of Stanford Bio-X, MCHRI and the Wu Tsai Neurosciences Institute , and a faculty fellow of Stanford ChEM-H .

To read all stories about Stanford science, subscribe to the biweekly  Stanford Science Digest .

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Taylor Kubota, Stanford News Service: (650) 724-7707;  [email protected]

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Evotourism ®

A Smithsonian magazine special report

AT THE SMITHSONIAN

Seven new things we learned about human evolution in 2021.

Paleoanthropologists Briana Pobiner and Ryan McRae reveal some of the year’s best findings in human origins studies

Briana Pobiner and Ryan McRae

This year—2021—has been a year of progress in overcoming the effects of the Covid-19 pandemic on human evolution research. With some research projects around the world back up and running, we wanted to highlight new and exciting discoveries from 13 different countries on five different continents. Human evolution is the study of what links us all together, and we hope you enjoy these stories we picked to show the geographic and cultural diversity of human evolution research, as well as the different types of evidence for human evolution, including fossils, archaeology, genetics, and even footprints!

New Paranthropus robustus fossils from South Africa show microevolution within a single species.

The human fossil record, like any fossil record, is full of gaps and incomplete specimens that make our understanding of complex evolutionary trends difficult. Identifying species and the process by which new species emerge from fossils falls in the realm of macroevolution , or evolution over broad time scales. These trends and changes tend to be more pronounced and easier to identify in the fossil record; think about how different a Tyrannosaurus rex and a saber-toothed cat are from each other. Human evolution only took place over the course of 5 to 8 million years, a much shorter span compared to the roughly 200 million years since dinosaurs and mammals shared a common ancestor. Because of this, smaller-scale evolutionary changes within a single species or lineage over time, called microevolution , is often difficult to detect.

Fossils of one early human species, Paranthropus robustus , are known from multiple cave sites in South Africa. Like other Paranthropus species, P. robustus is defined by large, broad cheeks, massive molars and premolars, and a skull highly adapted for intense chewing. Fossils of P. robustus from Swartkrans cave, just 20 miles west of Johannesburg, are dated to around 1.8 million years ago and show a distinct sagittal crest, or ridge of bone along the top of the skull, with their jaws indicating a more efficient bite force. Newly discovered fossils of P. robustus from Drimolen cave , about 25 miles north of Johannesburg, described by Jesse Martin from La Trobe University and colleagues in January, are at least 200,000 years older (2.04-1.95 million years old) and have a differently positioned sagittal crest and a less efficient bite force, among other small differences. Despite numerous disparities between fossils at the two sites, they much more closely resemble each other than any other known species of hominin. Because of this, researchers kept them as the same species from two different time points in a single lineage . The differences between fossils at the two sites highlight microevolution within this Paranthropus lineage .

Fossil children from Kenya, France, and South Africa tell us how ancient and modern human burial practices changed over time.

Most of the human fossil record includes the remains of adult individuals; that’s likely because larger and thicker adult bones, and bones of larger individuals, are more likely to survive the burial, fossilization, and discovery processes. The fossil record also gets much richer after the practice of intentional human burial began, starting at least 100,000 years ago .

In November, María Martinón-Torres from CENIEH (National Research Center on Human Evolution) in Spain, Nicole Boivin and Michael Petraglia from the Max Planck Institute for the Science of Human History in Germany, and other colleagues announced the oldest known human burial in Africa —a two-and-a-half to three-year-old child from the site of Panga ya Saidi in Kenya. The child, nicknamed “Mtoto” which means “child" in Kiswahili, was deliberately buried in a tightly flexed position about 78,000 years ago, according to luminescence dating. The way the child’s head was positioned indicates possible burial with a perishable support, like a pillow. In December, a team led by University of Colorado, Denver’s Jaime Hodgkins reported the oldest known burial of a female modern human infant in Europe . She was buried in Arma Veirana Cave in Italy 10,000 years ago with an eagle-owl talon, four shell pendants, and more than 60 shell beads with patterns of wear indicating that adults had clearly worn them for a long time beforehand. This evidence indicates her treatment as a full person by the Mesolithic hunter-gatherer group she belonged to. After extracted DNA determined that she was a girl, the team nicknamed her “Neve” which means “snow” in Italian. Aside from our own species, Neanderthals are also known to sometimes purposefully bury their dead . In December, a team led by Antoine Balzeau from the CNRS (the French National Centre for Scientific Research) and Muséum National d’Histoire Naturelle in France and Asier Gómez-Olivencia from the University of the Basque Country in Spain provided both new and re-studied information on the archaeological context of the La Ferrassie 8 Neanderthal skeleton, a two-year-old buried in France about 41,000 years ago. They conclude that this child, who is one of the most recently directly dated Neanderthals (by Carbon-14) and whose partial skeleton was originally excavated in 1970 and 1973, was purposefully buried . There have also been suggestions that a third species, Homo naledi , known from South Africa between about 335,000 and 236,000 years ago, purposefully buried their dead—though without any ritual context. In November, a team led by University of the Witwatersrand’s Lee Berger published two papers with details of skull and tooth fragments of a four to six-year-old Homo naledi child fossil , nicknamed “Leti” after the Setswana word “letimela” meaning “the lost one.” Given the location of the child’s skull found in a very narrow, remote and inaccessible part of the Rising Star cave system, about a half mile from Swartkrans, this first partial skull of a child of Homo naledi yet recovered might support the idea that this species also deliberately disposed of their dead.

The first Europeans had recent Neanderthal relatives, according to genetic evidence from Czechia and Bulgaria.

Modern humans, Homo sapiens , evolved in Africa and eventually made it to every corner of the world. That is not news. However, we are still understanding how and when the earliest human migrations occurred. We also know that our ancestors interacted with other species of humans at the time, including Neanderthals , based on genetic evidence of Neanderthal DNA in modern humans alive today—an average of 1.9 percent in Europeans.

Remains of some of the earliest humans in Europe were described this year by multiple teams, except they were not fully human. All three of the earliest Homo sapiens in Europe exhibit evidence of Neanderthal interbreeding (admixture) in their recent genealogical past. In April, Kay Prüfer and a team from the Max Planck Institute for the Science of Human History described a human skull from Zlatý kůň, Czechia, dating to around 45,000 years old . This skull contains roughly 3.2 percent Neanderthal DNA in the highly variable regions of the genome, comparable with other humans from around that time. Interestingly, some of these regions indicating Neanderthal admixture were not the same as modern humans, and this individual is not directly ancestral to any population of modern humans, meaning they belonged to a population that has no living descendants. Also in April, Mateja Hajdinjak and a team from the Max Planck Institute for Evolutionary Anthropology described three similar genomes from individuals found in Bacho Kiro Cave, Bulgaria, dating between 46,000 and 42,000 years old . These individuals carry 3.8, 3.4, and 3.0 percent Neanderthal DNA, more than the modern human average. Based on the distribution of these sequences, the team concluded that the three individuals each had a Neanderthal ancestor only six or seven generations back. This is roughly the equivalent length of time from the turn of the twentieth century to today. Interestingly, these three genomes represent two distinct populations of humans that occupied the Bulgarian cave—one of which is directly ancestral to east Asian populations and Indigenous Americans, the other of which is directly ancestral to later western Europeans. These findings suggest that there is continuity of human occupation of Eurasia from the earliest known individuals to present day and that mixing with Neanderthals was likely common, even among different Homo sapiens populations.

A warty pig from Indonesia, a kangaroo from Australia, and a conch shell instrument from France all represent different forms of ancient art.

Currently, the world’s oldest representational or figurative art is a cave painting of a Sulawesi warty pig found in Leang Tedongnge, Indonesia, that was dated to at least 45,500 years ago using Uranium series dating—and reported in January by a team led by Adam Brumm and Maxime Aubert from Griffith University. In February, a team led by Damien Finch from the University of Melbourne in Australia worked with the Balanggarra Aboriginal Corporation, which represents the Traditional Owners of the land in the Kimberly region of Australia, to radiocarbon date mud wasp nests from rock shelters in this area. While there is fossil evidence of modern humans in Australia dating back to at least 50,000 years ago , this team determined that the oldest known Australian Aboriginal figurative rock paintings date back to between around 17,000 and 13,000 years ago . The naturalistic rock paintings mainly depict animals and some plants; the oldest example is of a about 6.5 footlong kangaroo painting on the ceiling of a rock shelter dated to around 17,300 years ago. Right around that time, about 18,000 years ago, an ancient human in France cut off the top of a conch shell and trimmed its jagged outer lip smooth so it could be used as the world’s oldest wind instrument . A team led by Carole Fritz and Gilles Tostello from the Université de Toulouse in France reported in February that they re-examined this shell, discovered in Marsoulas Cave in 1931, using CT scanning. In addition to the modifications described above, they found red fingerprint-sized and shaped dots on the internal surface of the shell, made with ochre pigment also used to create art on the walls of the cave. They also found traces of a wax or resin around the broken opening, which they interpreted as traces of an adhesive used to attach a mouthpiece as found in other conch shell instruments.

Fossil finds from China and Israel complicate the landscape of human diversity in the late Pleistocene.

This year a new species was named from fossil material found in northeast China: Homo longi . A team from Hebei University in China including Qiang Ji, Xijun Ni, Qingfeng Shao and colleagues described this new species dating to at least 146,000 years old. The story behind the discovery of this cranium is fascinating! It was hidden in a well from the Japanese occupying forces in the town of Harbin for 80 years and only recently rediscovered. Due to this history, the dating and provenience of the cranium are difficult to ascertain, but the morphology suggests a mosaic of primitive-like features as seen in Homo heidelbergensis , and other more derived features as seen in Homo sapiens and Neanderthals . Although the cranium closely resembles some other east Asian finds such as the Dali cranium , the team named a new species based on the unique suite of features. This newly named species may represent a distinct new lineage, or may potentially be the first cranial evidence of an enigmatic group of recent human relatives—the Denisovans . Adding to the increasingly complex picture of late Pleistocene Homo are finds from Nesher Ramla in Israel dating to 120,000 to 130,000 years old , described in June by Tel Aviv University’s Israel Hershkovitz and colleagues. Like the Homo longi cranium, the parietal bone, mandible and teeth recovered from Nesher Ramla exhibit a mix of primitive and derived features. The parietal and mandible have stronger affiliations with archaic Homo , such as Homo erectus , while all three parts have features linking them to Neanderthals. Declining to name a new species , the team instead suggests that these finds may represent a link between earlier fossils with “Neanderthal-like features” from Qesem Cave and other sites around 400,000 years ago to later occupation by full Neanderthals closer to 70,000 years ago. Regardless of what these finds may come to represent in the form of new species, they tell us that modern-like traits did not evolve simultaneously, and that the landscape of human interaction in the late Pleistocene was more complex than we realize.

The ghosts of modern humans past were found in DNA in dirt from Denisova Cave in Russia.

Denisova Cave in Russia, which has yielded fossil evidence of Denisovans and Neanderthals (and even remains of a 13-year-old girl who was a hybrid with a Neanderthal mother and Denisovan father), is a paleoanthropological gift that keeps on giving! In June, a team led by Elena Zavala and Matthias Meyer from the Max Planck Institute for Evolutionary Anthropology in Germany and Zenobia Jacobs and Richard Roberts from the University of Wollongong in Australia analyzed DNA from 728 sediment samples from Denisova Cave —the largest analysis ever of sediment DNA from a single excavation site. They found ancient DNA from Denisovans and Neanderthals… and modern humans, whose fossils have not been found there, but who were suspected to have lived there based on Upper Paleolithic jewelry typically made by ancient modern humans found in 45,000-year-old layers there. The study also provided more details about the timing and environmental conditions of occupation of the cave by these three hominin species: first Denisovans were there, between 250,000 and 170,000 years ago; then Neanderthals arrived at the end of this time period (during a colder period) and joined the Denisovans, except between 130,000 and 100,000 years ago (during a warmer period) when only Neanderthal DNA was detected. The Denisovans who came back to the cave after 100,000 years ago have different mitochondrial DNA, suggesting they were from a different population. Finally, modern humans arrived at Denisova Cave by 45,000 years ago. Both fossil and genetic evidence point to a landscape of multiple interacting human species in the late Pleistocene, and it seems like Denisova Cave was the place to be!

Fossilized footprints bring to light new interpretations of behavior and migration in Tanzania, the United States and Spain.

Usually when we think of fossils, we think of the mineralized remnants of bone that represent the skeletons of long since passed organisms. Yet trace fossils, such as fossilized footprints, give us direct evidence of organisms at a specific place in a specific time. The Laetoli footprints , for example, represent the earliest undoubted bipedal hominin, Australopithecus afarensis (Lucy’s species) at 3.6 million years ago. In December, a team led by Ellison McNutt from Ohio University announced that their reanalysis of some of the footprints from Site A at Laetoli were not left by a bear, as had been hypothesized, but by a bipedal hominin. Furthermore, because they are so different from the well-known footprints from Site G, they represent a different bipedal species walking within 1 kilometer (0.6 miles) of each other within the span of a few days! Recently uncovered and dated footprints in White Sands National Park , New Mexico, described in September by a team led by Matthew Bennett of Bournemouth University, place modern humans in the area between 23,000 to 21,000 years ago. Hypotheses as to how Indigenous Americans migrated into North America vary in terms of method (ice-free land corridor versus coastal route) as well as timing. Regardless of the means by which people traveled to North America, migration was highly unlikely, if not impossible, during the last glacial maximum (LGM), roughly 26,000 to 20,000 years ago. These footprints place modern humans south of the ice sheet during this period, meaning that they most likely migrated prior to the LGM . This significantly expands the duration of human occupation past the 13,000 years ago supported by Clovis culture and the roughly 20,000 years ago supported by other evidence. Furthermore, it means that humans and megafauna, like giant ground sloths and wooly mammoths, coexisted for longer than previously thought, potentially lending credit to the theory that their extinction was not caused by humans. Also interesting is that most of these footprints were likely made by children and teenagers, potentially pointing to division of labor within a community. Speaking of footprints left by ancient children, a team led by Eduardo Mayoral from Universidad de Huelva reported 87 Neanderthal footprints from the seaside site of Matalascañas in southwestern Spain in March. Dated at about 106,000 years ago, these are now the oldest Neanderthal footprints in Europe, and possibly in the world. The researchers conclude that of the 36 Neanderthals that left these footprints, 11 were children; the group may have been hunting for birds and small animals, fishing, searching for shellfish… or just frolicking on the seashore. Aw.

A version of this article  was originally published  on the PLOS SciComm blog.

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Briana Pobiner | READ MORE

Briana Pobiner is a paleoanthropologist with the National Museum of Natural History’s Human Origins Program . She lead's the program's education and outreach efforts. 

Ryan McRae | READ MORE

Dr. Ryan McRae is a paleoanthropologist studying the hominin fossil record on a macroscopic scale. He currently works for the National Museum of Natural History’s Human Origins Program as a contractor focusing on research, education, and outreach, and is an adjunct assistant professor of anatomy at the George Washington University School of Medicine and Health Sciences.

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Research in human evolution is a strongly multi-disciplinary programme. The overarching framework is that understanding how humans evolved requires approaches from fields as diverse as human palaeontology, genomics, the cognitive sciences, behavioural ecology and archaeology. The Leverhulme Centre for Human Evolutionary studies at Cambridge is one of the leading research centres in the field, and encourages and supports this broad-based approach.

Human evolutionary studies at Cambridge are led by eight PIs, whose research covers a full  range of topics – the evolution of modern humans in Africa; behaviour, ecology and cognition of hunter-gatherers; the evolution of culture and technology in apes; health and disease in human evolution; growth and development in a life history context in contemporary populations, computational and bioinformatics approaches to human evolution and genetics; cultural evolution; comparative evolutionary ecology. In addition, there are close collaborations with other research groups in Archaeology (Palaeolithic Archaeology, Proteomics, Isotope Ecology), and in other Departments (Ancient DNA, Language Sciences)

There is a strong focus on Africa among the Cambridge human evolutionary studies researchers – Kenya, Congo, South Africa, Ivory Coast – but research is also carried out in Iraq, Bangladesh, Malaysia, New Guinea and within Europe. Research involves fieldwork, development and application of new scientific methodologies, and computational approaches.

What unites these multiple groups is a common interest in the application of evolutionary principles and methodologies to problems and datasets, to elucidate the patterns and process of human evolutionary history and how it impacts on current humanity.

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• ORIGINAL SCIENTIFIC ARTICLE
• Open access
• Published: 29 July 2010

Human Origins Studies: A Historical Perspective

• Tom Gundling 1  

Evolution: Education and Outreach volume  3 ,  pages 314–321 ( 2010 ) Cite this article

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Research into the deep history of the human species is a relatively young science which can be divided into two broad periods. The first spans the century between the publication of Darwin’s Origin and the end of World War II. This period is characterized by the recovery of the first non-modern human fossils and subsequent attempts at reconstructing family trees as visual representations of the transition from ape to human. The second period, from 1945 to the present, is marked by a dramatic upsurge in the quantity of research, with a concomitant increase in specialization. During this time, emphasis shifted from classification of fossil humans to paleoecology in which hominids were seen as parts of complex evolving ecosystems. This shift is in no small part due to the incorporation of neo-Darwinian synthetic theory. Finally, technological innovation and changes in social context are considered as influences on human origins studies.

Introduction

Considering the grand sweep of history, the realization that human beings gradually evolved from some non-human ancestor represents a very recent insight. Even so, the goal of this one brief essay cannot be to provide an in-depth description and analysis of every significant development within the field of paleoanthropology, but rather to identify broad patterns and highlight a collection of “events” that are most germane in shaping current understanding of our evolutionary origin. These events naturally include the accretion of fossil material, the raw data which is the direct, if mute, testimony of the past. These fossil discoveries are situated among technological breakthroughs, theoretical shifts, and changes in the sociocultural context in which human origins studies were conducted. It is only through such a contextualized historical approach that we can truly grasp our current understanding of human origins. Foibles of the past remind us to be critical in assessing newly produced knowledge, yet simultaneously we can genuinely appreciate the enormous strides that have been made.

In addition to selective coverage, a second caveat is that this review will focus on research by scientists writing in English. Non-modern hominids Footnote 1 are a cosmopolitan bunch, having been discovered throughout Africa, Asia, and Europe, and there is a significant literature in other languages. In an effort to ameliorate both of these shortcomings, numerous secondary references are included in the bibliography, providing more in-depth information on specific topics. For example, some texts approach the history of paleoanthropology by detailing a single time period (Bowler 1986 ), early human species (Walker and Shipman 1996 ), or researcher (Morell 1995 ), and there are quite a few that consider the subject more comprehensively (Leakey and Goodall 1969 ; Reader 1988 ; Lewin 1997 ; Tattersall 2008 ). In addition, there are a handful of encyclopedia format tomes (Jones et al. 1994 ; Spencer 1997 ; Delson et al. 2000 ), textbooks (Conroy 2005 ; Cela-Conde and Ayala 2007 ; Klein 2009 ), and “coffee table” popular volumes (Stringer and Andrews 2005 ; Johanson and Edgar 2006 ) that in part address the history of human origins studies. Moreover, these texts contain abundant references to the primary literature if that level of scrutiny is desired.

In seeking to provide a useful heuristic framework for the purposes of this particular essay, human origins studies can be broken down into two very broad periods. The first is roughly the century between 1850 and 1950 when research, often conducted by individuals with training outside of anthropology, focused on taxonomy and phylogeny. In other words, although scientists were cognizant that climate change (e.g., northern hemisphere glaciations) would have directly impacted the evolution of early humans, they were mainly interested in collecting “missing links,” naming them, and creating family trees. The second period, from 1950 to the present, is characterized by the relatively rapid development of paleoanthropology as it is currently practiced. Here the emphasis is only partly on the hominids themselves, with ecological context being of equal importance.

The Emergence of Human Origins Studies

This review begins with two mid-nineteenth century developments which are often conflated, but were initially distinct. The first is the acceptance of a temporal association of human material culture (stone tools), with extinct Ice Age mammals (Van Riper 1993 ; Sommer 2007 ). This was significant in that it opened up a considerable prehistory for the human species, well beyond estimates derived from literal scriptural interpretation. However, while acknowledging a lengthy antiquity for the human species, there was, at the time, no reason to suspect that the makers of the stone tools were not fully modern humans in a biological sense. The second major development was the publication of Darwin’s On the Origin of Species in 1859 (Darwin 1859 ). Darwin is rightfully credited with being the most influential, although by no means the first individual to broach the subject of descent with modification, or transmutation theory, as he put it (for an overview of pre-evolutionary ideas related to human origins, see Greene 1959 , Bowler 2003 ). Darwin’s central thesis was that all living species shared a common ancestry, with “endless forms most beautiful” having diverged via natural selection, and although he only briefly mentioned his own species the inference was clear. These two events dovetailed into the now quotidian, but then controversial, notion that humans had evolved over a vast expanse of time (Grayson 1983 ).

While Darwin was initially reticent to discuss human evolution in any detail, his colleague Thomas H. Huxley harbored no such reluctance when he published Man’s Place in Nature: Essays in 1863 (Huxley 1900 ). Darwin freely admitted that the veracity of his audacious proposal would have to withstand paleontological scrutiny and that his theory would collapse in the absence of transitional fossil forms. Huxley’s advantage, beyond his more outspoken personality, was that he actually had a fossil human to describe. The first Neandertal recognized by science was discovered in 1856; however, its description only appeared in English three years later, just as Darwin was going to press (Trinkaus and Shipman 1993 ). Huxley provided a detailed description of the eponymous cranium coupled with carefully composed line drawings (Huxley 1900 ). However, while the importance of the Neandertals in providing empirical evidence documenting an ancient and morphologically distinct human form cannot be discounted, these people hardly bridged the gap separating humans and the great apes. Although a few dissenting voices denied the close evolutionary relationship among humans and the “man-like” apes, and consequently an ape phase of human ancestry, most scientists accepted the overwhelming morphological and embryological evidence in support of just such a relationship. This acceptance was in no small part due to Huxley’s meticulous comparison of gorilla and human anatomy in which he concluded that the gorilla and its close relation, the chimpanzee, represented the nearest approach to humanity in nature.

If Neandertals were more or less human, then more distant, primitive “missing links” remained to be discovered. Just such fossils were recovered on the island of Java in the 1890s by Dutch physician Eugene Dubois, who had traveled to Indonesia as part of the army but with the express purpose of finding the remains of primitive humans (Shipman 2001 ). Java Man consisted of a skull cap, a femur, and a few isolated teeth which taken in combination suggested an early human with a much smaller cranial capacity relative to Neandertals or Homo sapiens (roughly 1,000 vs. 1,500 cubic centimeters), although the femur appeared modern. Dubois did not receive the universal accolades and acceptance he coveted, but his fossils bolstered the conventional wisdom at the time that humans first evolved somewhere in Asia.

During the early twentieth century, the early hominid fossil record grew significantly, if not exponentially, and evolution was widely accepted in scientific circles even while large segments of the lay public remained skeptical. Certainly, there were disagreements over whether natural selection was a sufficient evolutionary mechanism in itself (Bowler 1983 ), but the basic premise of biological change through time was affirmed. The recovery of additional Neandertal remains in Europe refuted lingering claims of pathology regarding the original Neander Valley specimen and solidified the interpretation that the latter was representative of a population of archaic humans occupying Ice Age Europe. Some Neandertal remains were interpreted as not only indicating intentional internment but also associated funerary ritual. The European fossil record was extended significantly with the recovery of a robust lower jaw from Mauer, Germany discovered in 1907.

In 1912 in England, heretofore devoid of non-modern hominid remains despite the prominence of several British scholars in human origins studies, the announcement of hominid fossils from Piltdown was warmly received locally, if with some incredulity abroad. Piltdown was significant since it reified the “brain first” hypothesis, in which primitive humans evolved a large brain before other key human traits evolved. Although a favorite of intelligent design creationism advocates, Piltdown is actually a beautiful example of the scientific method at work, whereby new evidence eventually calls into question prior interpretation, and in this case recognition of intentional fraud (Spencer 1990 ). It was, after all, a new relative dating method measuring the fluorine content of fossils that in 1953 exposed the non-contemporaneity of the jaw and skull. In any case, in the first decades of the twentieth century, a fairly simple human family tree was beginning to emerge (see McCown and Kennedy 1972 and especially Delisle 2007 for exceptions). Relatively small-brained Pithecanthropus led to the more capacious Neandertals and Piltdown, who in turn evolved into modern H. sapiens . Yet the truly ape-like human ancestors remained elusive.

Africa as the Cradle of Humanity

In 1921 a skull bearing superficial resemblance to European Neandertals was recovered as part of mining operations at a place called Broken Hill in Northern Rhodesia (now Zambia). Rhodesian Man marks the recovery of the first in a very long line of non-modern hominids from the African continent. A mere four years later, University of Witwatersrand anatomist Raymond Dart, Australian by birth and having been trained in England, published a brief paper describing the fossil skull of a juvenile “ape” discovered in a limestone quarry near Taung, South Africa. Dart identified certain features of the face, the teeth, the cranium, and the brain of Australopithecus africanus that foreshadowed those of H. sapiens and made the startling claim that what was essentially a bipedal ape signaled the beginning of the human lineage separate from the African great apes.

Initially, with only the one individual, and a juvenile at that, Dart found little support. His most ardent advocate, Scottish physician and paleontologist Robert Broom discovered additional fragmentary remains of the australopithecines, as they were then called, in other South African caves in the 1930s, but these were initially insufficient to sway opinion (Dart 1959 ). This was perhaps due to the near-simultaneous discovery of significant hominid remains from Zhoukoutien (Dragon Bone Hill) in China which quickly eclipsed whatever controversy the diminutive skull from Taung elicited, and despite Broom’s ongoing efforts. As was the case with Java Man, the more complete Chinese fossils fulfilled the expectations of many scientists who anticipated that earliest human ancestors evolved to the East. Comparative analysis of the Javanese and Chinese fossils revealed a great deal of similarity, and all of the fossils were ultimately subsumed in the species Homo erectus .

The Neo-Darwinian Synthesis and the New Physical Anthropology

For several disparate reasons, the decades following the end of World War II (WWII) rather quickly led to a science of paleoanthropology that is recognizably modern. One significant factor relevant in the U.S. if not everywhere, was the dramatic upsurge in enrollment at colleges and universities. The G.I. Bill and subsequent effects of Civil Rights legislation that greatly increased access to higher education meant that millions more students went to college and hence the expansion of existing campuses and programs and in some cases the appearance of entirely new colleges and universities Footnote 2 . As a result, greater numbers of faculty were required who could teach courses and supervise research in diverse academic programs, which in turn led to an attendant rise in the numbers of graduate students themselves who went on to secure positions at institutions of higher learning. Consequently, many disciplines experienced significant increases in research activity, including physical anthropology, and it is worth noting that this was the first generation of researchers whose formal training was in physical anthropology, not in some allied field such as anatomy or medicine. The dramatic rise in practitioners not only increased the knowledge base in terms of simple quantity, but specialization within the field also began to emerge.

A second crucial development that transformed human evolutionary studies was theoretical in nature. Changing ideas regarding the process of evolution had been fermenting and roiling in biology circles for several decades before they infiltrated the study of human origins. In essence, a consensus was reached among biologists ( sensu lato ) that Darwinian natural selection acting on variation arising from random mutation was a sufficient mechanism to explain evolutionary change. For anthropologists, although questions of taxonomy and phylogeny remained important, the intellectual fallout of the so-called neo-Darwinian synthesis led to the “New Physical Anthropology” in which early hominid fossils, rather than representative of some platonic archetype, were interpreted as unique members of variable populations. Focusing on evolution as a process effecting change in populations over time, in contrast to the comparatively myopic sorting of the resulting pattern , arguably represents the most significant theoretical shift in thinking about evolution since Darwin.

Given the comparative de-emphasis on iconic types, the bloated alpha taxonomy of the past was reduced to a mere handful of hominid species displaying previously under-appreciated within species variability. This great reduction in hominid names and consequent simplification of hominid family trees has led some modern scholars to lament what they see as a return to the bad old days of teleology and orthogenesis. Yet there can be little doubt that the “splitting” taxonomic philosophy of the past where almost every new specimen received a new species or quite frequently a new genus name was in dire need of revision.

Just as species types came under scrutiny, so did the concept of evolutionary grades which had up to this point made clear distinctions between the categories of ape and human. While this may have provided some welcome taxonomic clarity, it was artificial in that it ignored the evolutionary reality that at some point members of the human lineage were very ape-like. This realization, obvious in retrospect, led to the widespread acceptance of the South African australopithecines as human ancestors, and the important corollary that bipedalism preceded other distinctive human attributes (Gundling 2005 ).

In addition to increased research activity and theoretical shifts, by the early 1960s technological innovations for the first time permitted the creation of a reliable absolute timescale of human evolution. Comparative protein analysis demonstrated that the African apes were most similar genetically to H. sapiens , inferring their recent common ancestry to the exclusion of other apes and monkeys. Molecular clocks based upon mutation rates and calibrated by the fossil record suggested that this common ancestor lived as little as a few million years ago, although recent estimates put this ancestor at seven to five million years ago. Consequently, known early and middle Miocene ape species became suspect as purported human ancestors, since they preceded the split between the hominid and great ape lineages. Most notably this eventually led to the downfall of Ramapithecus , a Miocene ape genus once widely hailed as a very ancient and very primitive hominid Lewin ( 1997 ).

While molecular studies of living species effectively imposed a theoretical maximum on the age of the hominid lineage, the temporal framework of human origins was further clarified with the introduction of the new potassium argon (K-Ar) method of absolute dating. Louis and Mary Leakey had been scouring the fossil-bearing sediments in and around eastern Africa’s Great Rift Valley for decades when Mary discovered the skull of a robust australopithecine at Olduvai Gorge in 1959. Significantly, Zinjanthropus , the genus coined for the new skull, was discovered within sediments near the base of the Pleistocene Epoch. Volcanic minerals from associated strata were dated to approximately 1.75 million years ago using the K–Ar method, nearly double the age estimated from using other more crude means. This greatly expanded time range certainly bolstered claims for the australopithecines as human ancestors rather than extinct collateral cousins to the “true” human lineage, yet to be discovered.

As an aside, Louis Leakey’s interest in understanding the human past was not limited to the collection of fossils. Sherwood Washburn, a main architect of the new physical anthropology, along with Irven DeVore, conducted pioneering studies of savanna baboons, large-bodied, terrestrial, and highly social primates that served as living proxies for modeling early hominid behavioral ecology (Washburn and DeVore 1961 ). Leakey, on the other hand, took a more phylogenetically based approach and hired scholars to conduct research into the behavior of the great apes as a potential new data source informing hypotheses of early hominid behavior. Jane Goodall was the first, studying chimpanzee behavior at Gombe in Tanzania, then came Dianne Fossey who undertook a longitudinal study of mountain gorillas in Rwanda, and finally Birute Galdikas traveled to Indonesia to conduct field studies of the orangutan (see Kinzey 1987 and De Waal 2001 for more recent primate studies that explicitly address questions of human behavioral evolution).

The emergence of paleoanthropology as a truly multidisciplinary endeavor, concerned with a more holistic picture of our evolutionary past, was a logical extension of the post-WWII new physical anthropology which eschewed simple classification and promoted variable populations as the unit of study. Naturally, these hominid populations did not exist in a vacuum but were components of complex, evolving ecosystems. Hence, field work began to emphasize the collection of greater contextual data in an effort to reconstruct biological and physical environments in which these human ancestors existed and evolved. One of the first field projects to adopt this new approach was an international expedition centered around the Omo River Valley in southern Ethiopia, beginning in 1967. Remarkably, of the 50 papers collected in the resulting volume, only five primarily focus on the hominid remains themselves (Coppens et al. 1976 ).

Early Human Diet and Subsistence

One major aspect of early hominid ecology that occupied researchers engaged in such multidisciplinary efforts was subsistence, which has understandably been of great interest to paleoanthropologists, particularly after 1950 as scientists endeavored to contextualize the fossil remains of distant ancestors. What early humans ate, how food was acquired and processed, even how it was distributed among members of a social group, became viable questions. Throughout the 1950s and 1960s it was widely assumed that the social, cognitive, and technological skills associated with big-game hunting drove the evolution of the human species; in fact the allure of “Man the Hunter” is longstanding in Western thought (Cartmill 1993 ). Raymond Dart, as part of his second foray into human origins studies, proposed that Australopithecus had already developed a hunting strategy facilitated by a technology comprised of durable animal parts that he referred to as the osteodontokeratic (bone, tooth, horn) culture. This concept was enthusiastically embraced by writer Robert Ardrey, who published a series of four popular novels documenting the success of these “killer apes” in the context of a changing environment (e.g., Ardrey 1976 ). Research scientists were only slightly less enthusiastic in championing such ideas (Lee and DeVore 1968 ) which remain popular, if more nuanced today (Wrangham and Peterson 1996 ).

Mirroring changes in the broader society, by the early 1970s some anthropologists challenged the “Man the Hunter” hypothesis and developed an alternative that focused on the central role of women in child rearing and gathering of food resources (Dahlberg 1981 ). These studies used ethnographic data from extant food-foraging societies, the rarity of which injected a sense of urgency on the part of anthropologists. Not long after the “Women the Gatherer” model appeared as a second wave feminist rejoinder to the previously unquestioned authority of “Man the Hunter,” another group of researchers also began to question the big-game hunting scenario. Archeologists, geologists, and paleontologists began working on “site formation processes” to get a better understanding of how assemblages of fragmented animal bones and stone tools came to be commingled. Over the next few decades, often with recourse to modern ecosystems as analogs, one of the main conclusions drawn from the new science called taphonomy (=laws of burial) was the potential importance of scavenging. The association of “bones and stones” was no longer assumed to be the signature of hominid big-game hunting but instead interpreted as meals containing essential fat and protein scavenged by early humans. Perhaps even more disconcerting, some sites were reinterpreted as the remains of carnivore kills occasionally including early humans themselves (Brain 1981 ; Hart and Sussman 2008 ).

Here’s Lucy

If Mary and Louis Leakey’s discoveries at Olduvai put the Great Rift Valley on the map, during the 1970s eastern Africa was validated as the center of early hominid studies. The Leakey’s son Richard established himself on the east side of Lake Turkana in northern Kenya, where his expeditions uncovered a prolific cache of early hominid fossils, some of which corroborated the occasionally controversial claims made by his parents a decade earlier. Sediments around the lake yielded hominid fossils of robust australopithecines, early members of genus Homo , and an early African variant of Asian H. erectus , these days referred to as Homo ergaster (Leakey and Lewin 1978 ). The latter includes a mostly complete skeleton, KNM-WT15000, which has become iconic for the species (Walker and Shipman 1996 ).

Arguably the most significant fossil discovery of the 1970s was another partial skeleton, AL-288, from Hadar, Ethiopia, better known as Lucy (Johanson and Edey 1981 ). Here was a single individual represented by numerous skeletal elements, and although her morphology was generally similar to the “gracile” australopithecines of South Africa, she was even more primitive in some respects. Consequently her discoverers coined a new species name, Australopithecus afarensis that included not only the Hadar specimens but fossils collected by Mary Leakey’s expedition at Laetoli in Tanzania. The latter is renowned for its famous footprint trail preserved in solidified volcanic ash, imparting convincing evidence for bipedalism at 3.6 million years ago. Hadar is also replete with datable volcanic sediments, and Lucy’s status as the most primitive hominid was reinforced by firm radiometric dates which placed the fossils at greater than 3.0 million years ago, at the time astonishingly ancient.

One other significant event from the 1970s bears mentioning. Although the American Journal of Physical Anthropology was first published in 1918, it is perhaps surprising that a journal explicitly dedicated to the study of human evolution did not appear in the U.S. until 1972. Since then the Journal of Human Evolution has been the premier academic forum for publications related to human evolution, and in 1992, the Paleoanthropology Society was established, which organizes its own conference and publishes an online journal.

Modern Human Origins

The question of modern human origins has been debated for centuries, long predating paleoanthropology as a scientific discipline. One of the central issues, which became particularly evident as Renaissance and Enlightenment Europeans began to travel the globe on a regular basis, was how to explain the physical diversity of human populations. Two broad perspectives emerged, one which viewed all people as having a single origin and another which believed that supposedly distinct races had separate origins. The pre-Darwinian debates between so-called monogenists and polygenists were recast with the advent of an evolutionary paradigm in the mid-nineteenth century. Within this new theoretical context, monogenists believed that all living humans evolved from a common ancestor that was already H. sapiens , while the polygenists believed that the races had deeper roots and had descended from different non-modern ancestors (e.g., H. erectus or in a few instances different ape species). A major step towards resolving this debate came in 1987 with an analysis of living human mitochondrial DNA diversity which concluded that H. sapiens had a recent African origin. The discovery of essentially modern human fossils at the 160,000-year-old site of Herto, Ehtiopia, provides paleontological support for a recent African origin, and many subsequent genetic studies have supported this basic conclusion. However, the possibility of some gene flow between migrating early modern humans and local archaic populations remains plausible (compare Stringer and McKie 1996 and Wolpoff and Caspari 1998 , also see Relethford 2003 for a geneticist’s perspective).

Conclusion: Twenty-First Century Paleoanthropology

New fossil discoveries, technological innovations, theoretical advances, and social transformations will continue to inform knowledge of our deep past. Recovery of hominid fossils, some from previously unknown time periods and geographic locations, continues at a brisk rate. Many of the most significant recent discoveries are beginning to fill in the crucial African late Miocene time period during which our lineage ramified from that leading to the chimpanzee (Gibbons 2006 ). Of particular note, one of these fossils was discovered in Chad, quite a distance from established sites in the Great Rift Valley, challenging the long standing hypothesis that hominids evolved in the savanna grasslands of eastern Africa while the African ape ancestors remained sequestered in their tropical rainforest refugium. Moreover, botanical, faunal, and geological evidence associated with very early fossil hominids in Ethiopia and Kenya intimate a forested environment, a discovery that clearly constrains hypotheses explaining the success of the bipedal adaptation.

Other significant fossil discoveries from the early Pleistocene site of Dmanisi in the Republic of Georgia have energized discussion of the initial expansion of early humans beyond the tropics of Africa (Wong 2006 ). Not only are these fossils considerably older than prior known Eurasian specimens, but they are morphologically primitive, especially in terms of stature and cranial capacity, and are associated with very simple (“mode 1”) lithic technology. These early migrants hardly manifest the tall striding bipeds equipped with comparatively advanced Acheulian bifacial tools so often depicted in earlier “out of Africa” scenarios Footnote 3 , which are at least in part based on the iconic WT15000 skeleton mentioned earlier.

Perhaps the most surprising discovery of the last decade is the diminutive 18,000-year-old skeleton from the Indonesian island of Flores, which has sparked a spirited, occasionally acrimonious debate between those advocates of a replacement model of modern human origins and those inclined towards regional continuity (Morwood and van Oosterzee 2007 ). The former, comprised of the team who made the discovery and their allies, interpret the remains as those of a surprisingly primitive hominid akin to early Homo , and perhaps the first documented example of the effects of island dwarfing on an early human population. Other scholars believe the remains to be those of a pathological modern human, whose illness resulted in a cascade of skeletal and dental anomalies. Ongoing research on Flores and other nearby locations will undoubtedly resolve this debate.

New discoveries are not limited to the paleontological record but also include behavioral information gleaned from archaeology. Symbolic expression in the form of language, art (including music), and religion is undoubtedly one of the most distinctive human traits. Evidence for such behavior has proved elusive beyond the seeming cultural explosion perceived in the Upper Paleolithic of Europe beginning around 35,000 years before present. However, archeological evidence for at least some of these behaviors has recently been coaxed out of several sites in sub-Saharan Africa. Advanced utilitarian objects such as blades and harpoons have been recovered well back into the Middle Stone Age and use of ochre and shells for body adornment has been found at sites approaching 100 kiloannum (Balter 2009 ).

Recent advances also include a plethora of technological innovations that have allowed anthropologists to hone traditional inquiries in the areas of dating (e.g., single crystal, laser fusion, argon–argon dating), systematic analysis (e.g., geometric morphometrics), and paleoenvironmental reconstruction (e.g., stable isotope analysis). The badly distorted remains of the spectacular 4.4 megaanum skeleton of Ardipithecus ramidus from Aramis, Ethiopia was restored in part using digital imaging technology (Gibbons 2009 ). Additionally, new technology is facilitating, perhaps even driving, novel questions such as those related to the emergence of the unique human life history pattern.

While fossils provide real-time evidence for human evolution, signals from our ancient past are also encoded into our modern DNA. The groundbreaking work of the 1960s effectively demonstrated our close affinity with the African great apes, and today’s genomic analyses comparing humans and chimpanzees are beginning to reveal differences in much finer detail than heretofore possible. Already several areas within the human genome have been identified as having undergone intense selection; these regions may be related to the evolution of the especially dexterous human thumb, reduction of muscles of mastication in the wake of the ability to cook food, the greatly enlarged neo-cortex, and our ability for spoken language.

In addition to modern DNA analyses, ancient DNA analysis has informed the “Neandertal problem” providing preliminary evidence in support of the replacement hypothesis, at least in Europe, whereby modern humans arriving there equipped with Upper Paleolithic technology drove the indigenous Neandertals to extinction. Even more recent genomic analyses, however, suggest that a small but detectable degree of interbreeding occurred when expanding modern human populations emerging from the African tropics encountered Neandertal populations in the Middle East around 120,000 years before present (Gibbons 2010 ).

In conclusion, our understanding of human origins, like all scientific knowledge, is the result of an ongoing, iterative process. Over the last few decades, the accelerating pace of fossil discoveries and the incorporation of innovative technologies have corroborated and enhanced much of what we already suspected to be true, although there have been a few surprises. No doubt this pattern will continue into the foreseeable future as we slowly, yet inexorably, piece together the circumstances by which our lineage became human.

Hominidae (=hominid) is the biological group (clade) to which humans and their extinct ancestors belong. For many current scholars, this group is distinguished at some lower taxonomic level, usually the tribe Hominini (=hominin). In this study, I maintain the traditional use of Hominidae simply to be consistent with the historical literature. For the same reason, I use the subfamily designation Australopithecinae (=australopithecine) for all of the African “bipedal apes.” This group is certainly paraphyletic, to use the modern jargon, and as a result an increasing number of scholars prefer to use the less formal term australopith to lump together the various African species.

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I usually avoid this term despite its heavy usage within the scientific community. I believe that the “Out of Africa” trope perpetuates an anti-Africa bias which seems to suggest that early humans, on several occasions, left Africa wholesale as if there was something inherently undesirable about the place. I suppose that “hominid extra-tropical range expansion” doesn’t have the same ring, but it is more accurate.

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Biology and evolution of life science

1. introduction.

Biology literally means “the study of life”. Life Sciences attempts to untie the living things mysteries from the working of protein ‘machines’, to the growth of organism from a single cell to the majesty and intricacy of whole ecosystem. Questions about life sciences are as diverse and fascinating as life itself like; how a single cell knows to build up complex organism? How interpretation of genetic information takes place?

How the properties of organism are affected due to gene mutation? How ecosystem changes due to climate?

What can human genetic variation tell us about the history of human evolution and migration? Evolution is the change in heritable traits of biological populations over successive generations. Evolutionary processes give rise to diversity at every biological organization level. All life on earth shares a common ancestor known as the last universal ancestor. In the mid-19th century, Charles Darwin formulated the scientific theory of evolution by natural selection, while in the early 20th century the modern evolutionary synthesis integrated classical genetics with Darwin’s theory of evolution by natural selection through the discipline of population genetics. Evolution is a cornerstone of modern science, accepted as one of the most reliably established of all facts and theories of science, based on evidence not just from the biological sciences but also from anthropology, psychology, astrophysics, chemistry, geology, physics, mathematics, and other scientific disciplines, as well as behavioral and social sciences.

2. Theory of evolution on Earth

Today life diversity on earth is the result of evolution. On Earth life began at least 4 billion years ago and it has been evolving every year. In the beginning all living things on earth were single celled organism, after several years multicellular organism evolved after that diversity in life on earth increased day by day. Here in the figure shows the history of life on earth ( Fig. 1 ).

Timeline for history of life on Earth.

DNA (deoxyribonucleic acid) is the double helix structure shown in Fig. 2 . Its duplicate copies have coded information coiled up in almost all of the 100,000,000,000,000 (one hundred trillion) cells in your body. In human DNA has 46 segments; 23 segments received from father and 23 from mother. Each DNA contains exclusive information that determines what you look like, your personality and how your body cell is to function throughout your life.

Depictions of Saturn, DNA, and the Ark.

If one cell whole DNA was uncoiled and stretched out then it would be six feet long. Its detailed structure could not be seen due to its thin structure even under electron microscope. If all the coded information from one cell of one person were printed on books then it would fill a library of four thousand books and if the whole body DNA were positioned continuously, it would extend from here to Moon more than 500,000 times. If one set of DNA from each individual who still lived were placed in a pile, the final pile would weigh less than an aspirin.

3. Generic information

Different Scientists gave different information about genetic evolution like; Carl Sagan who showed by using simple calculation that how one cell’s value of genetic information approximates four thousand books of written information while volume of each book would have 50 cubic inches ( Sagan, 1977 ). 1014 cells are present in each adult individual. About 800 cubic miles have been worn from the Grand Canyon. According to that if each cell in one individual’s body was reduced to four thousand books then they would fill the Grand Canyon 98 times.

From earth the moon is 240,000 miles. If the human cell DNA were prolonged out and linked, it would be more than 7 feet long. If the entire DNA in one individual’s body were located back-to-back, it would enlarge to the moon 552,000 times.

The weight of DNA in human cell is 6.4 × 10 −12  g and almost less than fifty billion individuals lived on earth, if one copy of DNA from living individual were taken it is enough to define the physical characteristics of all those inhabitants in microscopic aspect and would weigh only, which is less than the weight of 1 aspirin.

According to Hoyle and Wickramasinghe, biochemical systems are exceptionally composite, so much so that the possibility of their being shaped from side to side haphazard shuffling of simple organic molecules is remarkably small, to a position certainly where it is inertly different from zero ( Hoyle and Wickramasinghe, 1999 ). Life cannot have a random beginning, like monkey’s troops thundering on typewriter could not be able to produce Shakespeare work. For the realistic cause entire visible universe is not vast adequate to hold the essential monkey hordes, essential typewriters, and surely the baskets for waste paper required for the deposition of wrong attempts. The same is true for the living matter.

The simple truth is not mentioned by Hoyle and Wickramasinghe that even a few correct words typed by monkey’s hordes would decompose long before a whole sentence of Shakespeare was completed. In the same way, a small number of correct amino acids sequences would decay long before a protein was completed, not to point out that thousands of proteins must be at their proper place in a living cell. At last the most composite condition of all is the occurrence of working DNA ( Vogel, 2001 ). They also state that our intelligence must reflect a vastly superior intelligence, even the tremendous idealized limit of God. They also believe that life was created by some intelligence somewhere in outer space and latter was transported to the Earth. All point mutations that have been studied on the molecular level turn out to reduce the genetic information and not to increase it ( Storz, 2002 ).

As Murray Eden reported that it is our contention that if ‘random’ is given a serious and crucial interpretation from a probabilistic opinion, then the randomness assumption is greatly improbable and a sufficient scientific theory of evolution has to wait for the finding and clarification of new natural laws like physical, physico-chemical, and biological ( Eden, 1967 ). I. After clearing up the above to a scientific symposium, Hoyle said that evolution was similar with the possibility that “a tornado sweeping through a junk-yard might assemble a Boeing 747 from the materials therein.

According to Ohno’s likable term is junk DNA that traps and no doubt dispirited a generation of researchers from studying the huge amount of important “junk” DNA that did not code for proteins ( Ohno, 1972 ). This study made an insightful point that if all the DNAs of human, mice and other organisms were useful then after so many mutations that build up in hundreds of millions of years then those species become extinct.

In different species non coding DNA differs more as compared to protein coding DNA. If we find a particular protein coding gene in human then we find nearly the same gene in mice and that rule just does not work for narrow elements. The biggest mistake in the history of molecular biology is the failure to recognize the importance of introns ( Mattick, 2003 ).

In transcription regulation, replication, RNA processing, translation and protein degradation non coding RNAs play an important role. Recent studies show that non coding RNAs are more important and abundant as compared to those initially imagined. The term junk DNA which is used is the reflection of our ignorance, non gene sequence also has their regulatory role ( Birney, 2012 ).

Fig. 3 shows that macroevolution would need a rising change in the complication of definite traits and organs while the microevolution is involved only in horizontal changes with no rising complications. Most of the creationists agree that natural selection occurs but it does not result in macroevolution.

Macroevolution vs. microevolution.

Today, the most accepted theory of life on Earth is evolution, and there is a vast amount of evidence supporting this theory. However, this was not always the case. Evolution can be described as a change in species over time. Dinosaur fossils are significant evidence of evolution and of past life on Earth. Before taking into consideration that how life began, first of all we understand the term organic evolution. It is naturally occurring and beneficial change that produces rising and inheritable complication. If the offspring of one form of life had a different and improved set of vital organs then this is called macroevolution, but the microevolution does not increase the complexity. By one or more mutation only size, shape and color are altered ( Taubes, 2009 ). Microevolution can be thought of as horizontal change, while macroevolution would involve vertical beneficial change in complexity. So the combination of microevolution and time will not produce macroevolution. Evolutionists have the same opinion that microevolution takes place. Since the start of history a minor change has been observed. But become aware of how frequently evolutionists give confirmation for microevolution to hold up macroevolution. It is macroevolution which requires new abilities and rising complication, resulting from new genetic information and is the center of the creation-evolution argument ( Maher, 2012 ).

4. The key parts of the theory of evolution

• • Charles Darwin’s observations and how they support the theory of evolution and the idea of natural selection.
• • The role of natural selection in adaptation.
• • Characteristics of micro evolutionary and macro evolutionary processes.

4.1. Top 5 misconceptions about evolution

4.1.1. it is just a theory.

In everyday language theory’ might mean a hunch or a guess. For scientists theory refers to a well supported explanation.

Scientific theories and scientific laws are often confused.

• * Evolution – The observation that organisms, including plants, bacteria and even molds change over time- depends on theory for explanation.
• * The most well know theory of evolution is the theory of natural selection.

4.1.2. Fittest survival

Is this accurate for Darwin’s theory of Natural Selection?

Fact 1 – Population tends to remain stable.

Fact 2 – Organisms reproduce more offspring than could be supported.

Interference 1 – Not all the offspring live long enough to reproduce.

Fact 3 – Resources are limited

Fact 4 – Individuals within population differ in individuality.

Fact 5 – Inherited characteristics are more.

Interference 2 – There will be differential survival and reproduction.

This is Natural Selection.

Interference 3 – Over time these differences will shift the makeup of the population.

This is decent with modification. Evolution will occur.

• * ‘Fit’ organisms will live & thrive to pass their genetic material to the next generation.
• * Fitness depends on reproducing & ensuring the survival of population rather than strength, speed or length.

4.1.3. Humans descend from Apes

• * Evolution holds that all life on Earth share common Ancestry.
• * Decent with modification means that human are unique as species, and we share many characteristics with other species.
• * Primates share 90% DNA sequence identity with humans.

4.1.4. No one was there and It cannot be Proven

• * Scientists operate like detectives.
• * With a few pieces of evidence about an event the investigator searches for clues that would legitimize or refute a claim.
• * Where is the support evidence coming from?

Evidence of evolution

• - Biochemistry (DNA).
• - Bones and fossils.
• - Comparative anatomy and physiology.
• - Computer modeling.
• - Modern experiments.
• - Developmental biology.

Journals publish evidence

• * Before publishing, a journal will send a manuscript to other scientists who review and critique it.
• * Peer review process often rejects manuscripts because there is not enough evidence to support the claims of the author. Science publishes less than 7% of submissions.
• * This level of organized skepticism is unique to Science.
• * Scientists become famous for overturning ideas and expanding paradigms.

4.1.5. Darwin was wrong

• * Darwin lived in a different time. He constructed the theory of Natural Selection from observing the finches in the Galapagos Islands and many other species across the world.
• * Genes was an unfamiliar term to that world.
• * Cells were seen but not manipulated.
• * Darwin’s mechanism continues to unify all biology – a contribution comparable to those of Newton or Einstein.
• * Today we define evaluation as changes in allelic frequency over time.
• * If we map different forms of genes (alleles) of a population and after a few generations the frequency changes, evolution has occurred.
• * This description is the best to date that captures the over-changing living world.
• * There are still many questions to ask and answer.
• * How do genes play a role in producing the features of organisms?
• * Why do mutations accumulate with different rates?
• * How do we protect our crops if pests evolve?
• * Evolution does not explain what started life, only how it persists, adapts, and changes.
• * Life need only begin once for evolution to occur.

4.2. The origins of life

In the process of evolution a series of natural changes cause species to arise, familiarize yourself to the environment, and turn out to be extinct.

Evolution   =   Change

By the process of biological evolution all species originated. The term species refers to a group who can reproduce their fertile offspring. Scientist classify the species with two scientific name first is genus name and second is species name like humans referred as Homo sapiens. In populations, there are variations or differences between individual members because of the variety of genes (alleles). Examples are skin color in humans, coat color in foxes. When there is a change in genes inherited from parents to offspring in different proportions then evolution occurs. These variations in genes arose for either (1) recombination of alleles when they sexually reproduce or (2) mutations.

Mechanism of evolution occurs by different ways

• 1. Natural selection.
• 2. Biased mutation.
• 3. Genetic drift.
• 4. Gene flow.

Recombining genetic material can happen in three ways.

• 1. Independent assortment.
• 2. Crossing over during meiosis.
• 3. Combining egg and sperm when fertilization occurs.

Mutations are usually neutral or harmful. Sometimes they can be beneficial if the environment is under a state of change.

• 1. Point mutation – In this there is change in a single base pair in DNA.
• 2. Frame shift – a single base pair is added or deleted from DNA.
• 3. Chromosome mutations – mistakes that affect the whole chromosome.
• 4. Deletion mutation – chromosome segments break off and do not reattach itself à new cell lacks genes carried by the segment that broke off.
• 5. Duplication or insertion mutation – Chromosome segments attach to a homologous chromosome that has lost the complementary segment. Result one chromosome carries two copies of one gene.
• 6. Inversion mutations – A segment of chromosome breaks off and then reattaches itself to the original chromosome backwards.
• 7. Translocation mutations – A chromosome segment attaches itself to a nonhomologous chromosome.

These variations lead to adaptations. Adaptations are traits that aid a population’s chance of survival and reproduction ( Hoyle, 1981 ).

A single individual does not change by the result of evolution, while it causes the change by inherited means of growth and development that are specified for a population. When the parent inherits these changes to the offspring then they become common in that population and as a result offspring inherit those genetic characteristics for probability of survival, capability to give birth which will work until the environment changes. Eventually, the genetic changes can modify a species overall way of life, like what it eats, how to grow, how it can live. As new genetic variations in early ancestor population’s preferential new abilities to become accustomed to environmental changes and so altered the human behavior causes the human evolution ( John, 2007 ).

5. Conclusion

Science should forever support conclusions on what is seen and reproducible. So what is observed? We see variations in lizard and birds. If macroevolution occurred in between forms they never as fossils.

An alert viewer can typically see astonishing discontinuities in these claimed upward changes, as well as in the drawing above. From the time of Darwin, different excuses made by evolutionists that why the world and our fossil museums are not spilling over with intermediates. Evolution is a scientific theory in biological sciences, which explains the emergence of new varieties of living things in the past and present. Evolution accounts for the conspicuous patterns of similarities and differences among living things over time and across habitats through the action of biological processes such as mutation, natural selection, symbiosis and genetic drift. Evolution has been subjected to scientific testing for over a century and has been again and again confirmed from different fields.

Peer review under responsibility of King Saud University.

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JSmol Viewer

Research progress on the application of one-step fabrication techniques for iridium-based thin films in the oxygen evolution reaction.

1. Introduction

2. catalytic mechanism of iridium oxide in oer, 3. one-step fabrication techniques for iridium-based thin films, 3.1. electrochemical deposition method, 3.2. physical vapor deposition method, 3.3. chemical vapor deposition method, 3.4. sol–gel method, 3.5. other methods, 4. future prospects, author contributions, institutional review board statement, informed consent statement, data availability statement, conflicts of interest.

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Li, W.; Zhu, J.; Cai, H.; Tong, Z.; Wang, X.; Wei, Y.; Wang, X.; Hu, C.; Zhao, X.; Zhang, X. Research Progress on the Application of One-Step Fabrication Techniques for Iridium-Based Thin Films in the Oxygen Evolution Reaction. Coatings 2024 , 14 , 1147. https://doi.org/10.3390/coatings14091147

Li W, Zhu J, Cai H, Tong Z, Wang X, Wei Y, Wang X, Hu C, Zhao X, Zhang X. Research Progress on the Application of One-Step Fabrication Techniques for Iridium-Based Thin Films in the Oxygen Evolution Reaction. Coatings . 2024; 14(9):1147. https://doi.org/10.3390/coatings14091147

Li, Wenting, Junyu Zhu, Hongzhong Cai, Zhongqiu Tong, Xian Wang, Yan Wei, Xingqiang Wang, Changyi Hu, Xingdong Zhao, and Xuxiang Zhang. 2024. "Research Progress on the Application of One-Step Fabrication Techniques for Iridium-Based Thin Films in the Oxygen Evolution Reaction" Coatings 14, no. 9: 1147. https://doi.org/10.3390/coatings14091147

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An Ellulian analysis of propaganda in the context of generative AI

• Original Paper
• Published: 04 September 2024
• Volume 26 , article number  60 , ( 2024 )

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• Xiaomei Bi 1 ,
• Xingyuan Su 2 &
• Xiaoyan Liu   ORCID: orcid.org/0009-0007-0342-8263 2  

The application of generative artificial intelligence (GenAI) technologies in the field of propaganda influences information creation, dissemination, and reception, and introduces new ethical challenges. This paper revisits the philosophical discourses of Jacques Ellul on technology and propaganda, placing them within the context of the rise of today’s generative AI technologies. Ellul identified the First Industrial Revolution as the initial juncture in the history of human technology that formed technique as a social phenomenon, which subsequently shaped the nature of propaganda as a technique. Subsequent developments in computer technology in the latter half of the 20th century enabled the formation of a technological system. This raises the question: Could generative AI represent another pivotal moment in the evolution of the technological system, and what are the ethical implications of propaganda technology in this context? This article seeks to illuminate current discussions on GenAI technology and propaganda ethics with Ellul’s insightful theoretical insights. In terms of research methodology, this study relies on textual interpretation and classical hermeneutics, including three processes: syntactic text interpretation, historical background, and situational application. Ellul’s research delves into the intrinsic links and inherent ethical dimensions between propaganda and technology, examining their comprehensive and enduring impacts. This normative perspective is crucial for a deep understanding of contemporary propaganda within the framework of emerging technologies, helping us to transcend the escalating spiral of propaganda technology and counter-propaganda techniques. By incorporating propaganda facilitated by generative AI technologies into the overall development logic of the technological society, this approach explores its ethical implications from a more macroscopic and holistic perspective.

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This study was supported by Shanghai Planning Office of Philosophy and Social Science (Project Title: Research on the Reform of China’s Cultural System in the New Era; NO. 2022ZDS002).

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The Invention of Prehistory: Empires, Violence, and Our Obsession with Human Origins

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The study of the human past matters for an array of positive and negative reasons. A new book from historian Stefanos Geroulanos promises to explore those reasons — but also to question the worth of studying human origins altogether — by reviewing the histories of the European–British–American quest to study human evolution, its allied projects, and its often racist, sexist and colonialist products. Geroulanos focuses on some of the more popular and political takes on this topic and the impacts and actions of some key scholars and popularizers (interestingly, including Sigmund Freud). Notably, he examines the sociopolitical implications of select aspects of human evolution scholarship, popularizing and politics across the late 19th and 20th centuries. In this respect, the book is a solid contribution and gets a lot right about the endeavour across the past few centuries. It would all be fine if Geroulanos intended this to be only a particular history of the subject. But he does not. Instead, he makes a more specific assertion: that the very “concept of humanity that follows from [the study of] prehistory rests on hierarchies and exclusions, on ideals of purity and power” (page 399). As a researcher of the field of human evolution today, I find its characterization incomplete and uninformed.

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