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The evolution of our understanding of human development over the last 10 years

Ali h. brivanlou.

1 Stem Cell Biology and Molecular Embryology Laboratory, The Rockefeller University, New York, NY USA

Norbert Gleicher

2 The Center for Human Reproduction, New York, NY USA

3 The Foundation for Reproductive Medicine, New York, NY USA

4 Department of Obstetrics and Gynecology, Medical University of Vienna, Vienna, Austria

As it fulfills an irresistible need to understand our own origins, research on human development occupies a unique niche in scientific and medical research. In this Comment, we explore the progress in our understanding of human development over the past 10 years. The focus is on basic research, clinical applications, and ethical considerations.

What basic research has taught us about human development

Over the last decade, progress in understanding our own development was mostly driven by the emergence and combination of remarkable new technologies. New molecular biology tools such as single-cell RNA-sequencing (sc-RNA-seq) unveiled the earliest genetic signature of the three cell lineages of the human blastocyst and allowed for the discovery of human-specific signatures 1 – 3 . CRISPR/Cas9 genome editing has offered further access to in vitro functional studies in human blastocysts 4 . However, as we discuss below, an ethical line was crossed when a group claimed that genetically modified human embryos had been transferred, leading to births 5 when neither public opinion nor a consensus within the scientific community had been reached regarding whether crossing the germline in in vitro fertilization (IVF) was safe and ethically acceptable.

On the embryology side, the development of an in vitro attachment platform for human blastocysts offered a first glance into post-implantation events up to 12 days 1 , 3 , 5 , 6 . This paved the way for several important discoveries, including the observation that the human embryo can self-organize to generate embryonic and extraembryonic germ layers, yolk sac, and amniotic cavities in the absence of maternal influences 5 , 6 ; and the presence of a transient embryonic tissue of trophectodermal lineage, adjacent to the yolk sac, therefore named, yolk-sac trophectoderm ( ysTE ) 5 . The presence of these seemingly human-specific populations was independently confirmed by sc-RNA-seq 1 .

The marriage of stem cell biology with bioengineering gave birth to the field of synthetic embryology 7 – 13 . This technology uses human embryonic stem cells (hESCs) cultured on geometrically confined micropatterned substrates to generate 2D in vitro models of human conceptuses, such as models of the gastrula ( gastruloids ) 7 – 13 , or parts of the embryo, such as cerebroids and neuruloids 14 . Thousands of nearly identical self-organizing human embryonic structures allow for standardization and reproducibility, which cannot be achieved in standard organoid structures 15 . Cells within these structures can be tracked and quantified in real time with sub-cellular resolution, using sophisticated quantification code, including artificial intelligence 14 .

Human gastruloids induce formation of the primitive streak and have enabled the deciphering of the molecular network underlying gastrulation—the most crucial moment of our lives 7 – 13 . 3D models of human epiblasts can spontaneously break axial symmetry, thus providing an assay for the elucidation of molecular events underlying the emergence of antero–posterior polarity 11 , 16 . A highly homogenous population of self-organizing 3D models of amniotic ectoderm-like cells can be obtained by combining microfluidic and microculture approaches 17 .

Finally, the development of interspecies chimeras provided the most stringent in vivo validation of human embryo models 9 , 10 , 18 . Unimaginable in human models, inter-species chimeras have become the next best choice to test whether hESC behavior in self-organizing gastruloids , as observed on microchips, would also occur in an embryonic environment 10 , 18 , 19 . Human/bird chimeras generated from transplanting human gastruloids into early chick embryos in ovo unexpectedly proved more efficient than previous methods 9 , 19 . They allowed for the observation of an entire self-organizing embryonic axis in bird eggs 9 . As birds are closer to dinosaurs than to humans, this high rate of success with these chimeras further suggested that these early patterning events must be highly conserved.

Translational clinical applications that arose from basic research

The past 10 years bore witness to significant clinical progress in reproductive medicine, often translated from basic research. Successful human uterus transplantation and the subsequent birth of healthy offspring was, for example, only achieved after years of meticulous laboratory work in animals 10 . Significant improvements in cryopreservation technology for human eggs and ovarian tissue were also preceded by research in model systems 10 , 20 . Practical clinical applications have been developed for women in need of cancer treatment that are toxic to ovaries. In these cases, oocytes and/or ovarian tissue can be cryopreserved for later use in fertility treatments once the patient is cured of her cancer 21 . This ever-evolving technology has already proven to result in live births, and has also become an integral part of routine infertility treatments with IVF, giving rise to the brand-new concept of fertility extension through egg-freezing.

Diagnostic technologies to assess retrieved eggs and preimplantation-stage embryos in the IVF process have been disappointing. For example, tracking extended embryo culture to blastocyst-stage with time-lapse imaging failed to improve embryo selection 22 . That chromosomal-abnormal embryos increase with maternal (but not paternal) age has been interpreted to mean that chromosomal abnormalities were a principal cause for lower implantation chances and higher miscarriage risks among older women. This assumption led to the rapidly growing utilization of chromosomal testing of human embryos prior to embryo transfer in a procedure recently renamed preimplantation genetic testing for aneuploidy (PGT-A) 23 . The hypothesis behind PGT-A is to exclude chromosomal-abnormal embryos from the transfer, thereby improving implantation potentials of remaining euploid embryos.

Here too, clinical evidence was unable to confirm the hypothesis 24 . Moreover, basic research demonstrated a self-correction mechanism in mouse 25 and human embryos 26 – 29 that arose during embryogenesis that was cell lineage-specific to the embryonic cell lineage. In contrast, PGT-A biopsies are obtained from the extraembryonic-derived trophectoderm, rendering any diagnostic procedure at the blastocyst stage ineffective. In addition, mathematical modeling demonstrated that results from a single trophectoderm biopsy could not be extrapolated to the whole embryo 30 . Transfer of PGT-A “chromosomal-abnormal diagnosed embryos” has resulted in the births of over 400 chromosomal-normal offspring 20 , 21 .

In recent years, increasing attention has also been given to the quickly evolving understanding of how interdependent lifestyle and human fertility are 31 – 33 , including the influence of diet on the microbiome, as in many other areas of medicine.

The ethical significance of understanding human development

Whether in clinical medicine or in the research laboratory, human embryology has remained an ethical minefield, strongly influenced by socio-political and religious considerations. At the core of the controversy resides the special moral value of the human embryo, a subject that has come to the forefront again with the ascent of human embryonic stem cell research 34 . There is, however, little consensus as to how to answer a previously raised question: “ what is an embryo ?” 35 . The term pre-embryo, first introduced in 1986, was defined as the interval up to the appearance of the primitive streak, which marks biological individuation at ~14 days post-fertilization. This definition designated the period beyond 14 days as the time when a pre-embryo attains special moral status 36 , 37 . Paradoxically, the term pre-embryo has been replaced by the indiscriminate use of the term embryo, whether at preimplantation cleavage or blastocyst-stages or post-implantation before day 14. It was suggested that the distinction was important for ethical, moral, and biological relevance. The principal reason is simple: Until a pre-embryo becomes an embryo, there is no way of knowing whether implantation has taken place, whether a pregnancy is developing, whether there is a single pregnancy or twinning, or whether fertilization ended up in a benign (hydatidiform mole) or even in a malignant tumor (choriocarcinoma) 35 . Assigning advanced moral value to embryos at those early stages is, therefore, difficult to defend.

The past 10 years have witnessed innumerous ethical debates related to this subject, each with its own social, historical, and religious justifications, reflecting cultural diversities in human populations. Most are triggered by scientific breakthroughs. We summarize here the major ethical challenges preoccupying reproductive research and clinical practice.

We have already briefly referred to CRISPR/Cas9 genome editing. While the use of sc-RNA-seq to identify the molecular blueprint of human development has not elicited significant controversy, CRISPR/Cas9 genome editing of human embryos has been a topic of intense discussions and is currently permissible only in vitro 38 . An alleged attempt in China of implanting human genome-edited embryos into the uterus supposedly led to two births (one a twin birth). Though widely discussed in the media, neither attempt was published in the medical literature, and therefore cannot be verified 5 , 38 .

The ethical debates surrounding the 14-day rule, quiescent since the early IVF days, experienced a rebirth that was prompted by in vitro human embryo attachment studies and the emergence of synthetic human embryos. Within this context, we note that self-organizing embryo models are nothing more than cells in culture and are certainly not embryos. Regardless of scientific merits, in the U.S., the National Institutes of Health (NIH) currently prohibits the use of public funds for the study of synthetic embryos “for ethical reasons”. After being under an NIH moratorium for more than a year, research on chimeras is now, however, again permitted, though human/non-human primate chimeras remain prohibited.

These ongoing ethical debates mostly also mirror those surrounding the lack of U.S. federal funding for clinical IVF and related research, as well as hESCs-derived model embryos. In this context, the American Society for Reproductive Medicine (ASRM)’s Ethics in Embryo Research Task Force recently made an important statement: “ Scientific research using human embryos advances human health and provides vital insights into reproduction and disease ” 39 .

Provided certain guidelines and safeguards are followed, research with already existing embryos or embryos specifically produced for research should be ethically acceptable as a means of obtaining new knowledge that may benefit human health. ASRM also pointed out that scientists and society must understand which research questions necessitate the use of human embryos.

It is gratifying to acknowledge the history and vitality of ongoing debates, especially since they increasingly mimic decision-making processes in the medical field. These debates are meant to be based on cost-benefit and/or risk-benefit assessments. These debates will, unquestionably, continue and, indeed, considering that every intervention has consequences, must be decided based on careful considerations, including all relevant stakeholders and all parts of society.

Acknowledgements

We like to thank Min Yang, Jean Marx Santel, Adam Souza, and Amir Brivanlou, for data gathering and critical reading of the manuscript, and constructive criticism.

Author contributions

Both A.H.B. and N.G. have contributed to writing the manuscript.

Competing interests

A.H.B. and N.G. are co-founders of OvaNova Inc. A.H.B. is a co-founder of Rumi Scientific Inc.

Peer review information Nature Communications thanks Alfonso Martinez Arias, Annelien Bredenoord and the other anonymous reviewer(s) for their contribution to the peer review of this work.

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

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• Published: 29 July 2021

The evolution of our understanding of human development over the last 10 years

• Ali H. Brivanlou   ORCID: orcid.org/0000-0002-1761-280X 1 &
• Norbert Gleicher   ORCID: orcid.org/0000-0002-0202-4167 2 , 3 , 4  

Nature Communications volume  12 , Article number:  4615 ( 2021 ) Cite this article

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• Developmental biology
• Embryogenesis

As it fulfills an irresistible need to understand our own origins, research on human development occupies a unique niche in scientific and medical research. In this Comment, we explore the progress in our understanding of human development over the past 10 years. The focus is on basic research, clinical applications, and ethical considerations.

What basic research has taught us about human development

Over the last decade, progress in understanding our own development was mostly driven by the emergence and combination of remarkable new technologies. New molecular biology tools such as single-cell RNA-sequencing (sc-RNA-seq) unveiled the earliest genetic signature of the three cell lineages of the human blastocyst and allowed for the discovery of human-specific signatures 1 , 2 , 3 . CRISPR/Cas9 genome editing has offered further access to in vitro functional studies in human blastocysts 4 . However, as we discuss below, an ethical line was crossed when a group claimed that genetically modified human embryos had been transferred, leading to births 5 when neither public opinion nor a consensus within the scientific community had been reached regarding whether crossing the germline in in vitro fertilization (IVF) was safe and ethically acceptable.

On the embryology side, the development of an in vitro attachment platform for human blastocysts offered a first glance into post-implantation events up to 12 days 1 , 3 , 5 , 6 . This paved the way for several important discoveries, including the observation that the human embryo can self-organize to generate embryonic and extraembryonic germ layers, yolk sac, and amniotic cavities in the absence of maternal influences 5 , 6 ; and the presence of a transient embryonic tissue of trophectodermal lineage, adjacent to the yolk sac, therefore named, yolk-sac trophectoderm ( ysTE ) 5 . The presence of these seemingly human-specific populations was independently confirmed by sc-RNA-seq 1 .

The marriage of stem cell biology with bioengineering gave birth to the field of synthetic embryology 7 , 8 , 9 , 10 , 11 , 12 , 13 . This technology uses human embryonic stem cells (hESCs) cultured on geometrically confined micropatterned substrates to generate 2D in vitro models of human conceptuses, such as models of the gastrula ( gastruloids ) 7 , 8 , 9 , 10 , 11 , 12 , 13 , or parts of the embryo, such as cerebroids and neuruloids 14 . Thousands of nearly identical self-organizing human embryonic structures allow for standardization and reproducibility, which cannot be achieved in standard organoid structures 15 . Cells within these structures can be tracked and quantified in real time with sub-cellular resolution, using sophisticated quantification code, including artificial intelligence 14 .

Human gastruloids induce formation of the primitive streak and have enabled the deciphering of the molecular network underlying gastrulation—the most crucial moment of our lives 7 , 8 , 9 , 10 , 11 , 12 , 13 . 3D models of human epiblasts can spontaneously break axial symmetry, thus providing an assay for the elucidation of molecular events underlying the emergence of antero–posterior polarity 11 , 16 . A highly homogenous population of self-organizing 3D models of amniotic ectoderm-like cells can be obtained by combining microfluidic and microculture approaches 17 .

Finally, the development of interspecies chimeras provided the most stringent in vivo validation of human embryo models 9 , 10 , 18 . Unimaginable in human models, inter-species chimeras have become the next best choice to test whether hESC behavior in self-organizing gastruloids , as observed on microchips, would also occur in an embryonic environment 10 , 18 , 19 . Human/bird chimeras generated from transplanting human gastruloids into early chick embryos in ovo unexpectedly proved more efficient than previous methods 9 , 19 . They allowed for the observation of an entire self-organizing embryonic axis in bird eggs 9 . As birds are closer to dinosaurs than to humans, this high rate of success with these chimeras further suggested that these early patterning events must be highly conserved.

Translational clinical applications that arose from basic research

The past 10 years bore witness to significant clinical progress in reproductive medicine, often translated from basic research. Successful human uterus transplantation and the subsequent birth of healthy offspring was, for example, only achieved after years of meticulous laboratory work in animals 10 . Significant improvements in cryopreservation technology for human eggs and ovarian tissue were also preceded by research in model systems 10 , 20 . Practical clinical applications have been developed for women in need of cancer treatment that are toxic to ovaries. In these cases, oocytes and/or ovarian tissue can be cryopreserved for later use in fertility treatments once the patient is cured of her cancer 21 . This ever-evolving technology has already proven to result in live births, and has also become an integral part of routine infertility treatments with IVF, giving rise to the brand-new concept of fertility extension through egg-freezing.

Diagnostic technologies to assess retrieved eggs and preimplantation-stage embryos in the IVF process have been disappointing. For example, tracking extended embryo culture to blastocyst-stage with time-lapse imaging failed to improve embryo selection 22 . That chromosomal-abnormal embryos increase with maternal (but not paternal) age has been interpreted to mean that chromosomal abnormalities were a principal cause for lower implantation chances and higher miscarriage risks among older women. This assumption led to the rapidly growing utilization of chromosomal testing of human embryos prior to embryo transfer in a procedure recently renamed preimplantation genetic testing for aneuploidy (PGT-A) 23 . The hypothesis behind PGT-A is to exclude chromosomal-abnormal embryos from the transfer, thereby improving implantation potentials of remaining euploid embryos.

Here too, clinical evidence was unable to confirm the hypothesis 24 . Moreover, basic research demonstrated a self-correction mechanism in mouse 25 and human embryos 26 , 27 , 28 , 29 that arose during embryogenesis that was cell lineage-specific to the embryonic cell lineage. In contrast, PGT-A biopsies are obtained from the extraembryonic-derived trophectoderm, rendering any diagnostic procedure at the blastocyst stage ineffective. In addition, mathematical modeling demonstrated that results from a single trophectoderm biopsy could not be extrapolated to the whole embryo 30 . Transfer of PGT-A “chromosomal-abnormal diagnosed embryos” has resulted in the births of over 400 chromosomal-normal offspring 20 , 21 .

In recent years, increasing attention has also been given to the quickly evolving understanding of how interdependent lifestyle and human fertility are 31 , 32 , 33 , including the influence of diet on the microbiome, as in many other areas of medicine.

The ethical significance of understanding human development

Whether in clinical medicine or in the research laboratory, human embryology has remained an ethical minefield, strongly influenced by socio-political and religious considerations. At the core of the controversy resides the special moral value of the human embryo, a subject that has come to the forefront again with the ascent of human embryonic stem cell research 34 . There is, however, little consensus as to how to answer a previously raised question: “ what is an embryo ?” 35 . The term pre-embryo, first introduced in 1986, was defined as the interval up to the appearance of the primitive streak, which marks biological individuation at ~14 days post-fertilization. This definition designated the period beyond 14 days as the time when a pre-embryo attains special moral status 36 , 37 . Paradoxically, the term pre-embryo has been replaced by the indiscriminate use of the term embryo, whether at preimplantation cleavage or blastocyst-stages or post-implantation before day 14. It was suggested that the distinction was important for ethical, moral, and biological relevance. The principal reason is simple: Until a pre-embryo becomes an embryo, there is no way of knowing whether implantation has taken place, whether a pregnancy is developing, whether there is a single pregnancy or twinning, or whether fertilization ended up in a benign (hydatidiform mole) or even in a malignant tumor (choriocarcinoma) 35 . Assigning advanced moral value to embryos at those early stages is, therefore, difficult to defend.

The past 10 years have witnessed innumerous ethical debates related to this subject, each with its own social, historical, and religious justifications, reflecting cultural diversities in human populations. Most are triggered by scientific breakthroughs. We summarize here the major ethical challenges preoccupying reproductive research and clinical practice.

We have already briefly referred to CRISPR/Cas9 genome editing. While the use of sc-RNA-seq to identify the molecular blueprint of human development has not elicited significant controversy, CRISPR/Cas9 genome editing of human embryos has been a topic of intense discussions and is currently permissible only in vitro 38 . An alleged attempt in China of implanting human genome-edited embryos into the uterus supposedly led to two births (one a twin birth). Though widely discussed in the media, neither attempt was published in the medical literature, and therefore cannot be verified 5 , 38 .

The ethical debates surrounding the 14-day rule, quiescent since the early IVF days, experienced a rebirth that was prompted by in vitro human embryo attachment studies and the emergence of synthetic human embryos. Within this context, we note that self-organizing embryo models are nothing more than cells in culture and are certainly not embryos. Regardless of scientific merits, in the U.S., the National Institutes of Health (NIH) currently prohibits the use of public funds for the study of synthetic embryos “for ethical reasons”. After being under an NIH moratorium for more than a year, research on chimeras is now, however, again permitted, though human/non-human primate chimeras remain prohibited.

These ongoing ethical debates mostly also mirror those surrounding the lack of U.S. federal funding for clinical IVF and related research, as well as hESCs-derived model embryos. In this context, the American Society for Reproductive Medicine (ASRM)’s Ethics in Embryo Research Task Force recently made an important statement: “ Scientific research using human embryos advances human health and provides vital insights into reproduction and disease ” 39 .

Provided certain guidelines and safeguards are followed, research with already existing embryos or embryos specifically produced for research should be ethically acceptable as a means of obtaining new knowledge that may benefit human health. ASRM also pointed out that scientists and society must understand which research questions necessitate the use of human embryos.

It is gratifying to acknowledge the history and vitality of ongoing debates, especially since they increasingly mimic decision-making processes in the medical field. These debates are meant to be based on cost-benefit and/or risk-benefit assessments. These debates will, unquestionably, continue and, indeed, considering that every intervention has consequences, must be decided based on careful considerations, including all relevant stakeholders and all parts of society.

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Acknowledgements

We like to thank Min Yang, Jean Marx Santel, Adam Souza, and Amir Brivanlou, for data gathering and critical reading of the manuscript, and constructive criticism.

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Ali H. Brivanlou

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A.H.B. and N.G. are co-founders of OvaNova Inc. A.H.B. is a co-founder of Rumi Scientific Inc.

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Peer review information Nature Communications thanks Alfonso Martinez Arias, Annelien Bredenoord and the other anonymous reviewer(s) for their contribution to the peer review of this work.

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Brivanlou, A.H., Gleicher, N. The evolution of our understanding of human development over the last 10 years. Nat Commun 12 , 4615 (2021). https://doi.org/10.1038/s41467-021-24793-3

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In September 2014, Development organised a four-day workshop titled ‘From Stem Cells to Human Development’. In planning this meeting, we sought to fill what we saw as a gap in the meeting calendar – a way of bringing together a diverse cross-section of researchers with a common interest in using the rapidly developing tools of stem cell biology, genetic engineering and genomic analysis to understand human development (for a review of the meeting, see Medvinsky and Livesey, 2015 ). The enthusiasm with which this workshop was met, from invited speakers and registered participants alike, confirmed our view that we are now in a period in which significant inroads into understanding human development will be made. With our ever-improving ability to model tissue development in vitro and to manipulate the human genome (and epigenome), we are now in a position to analyse human organogenesis and to understand how it differs from that in other model organisms – and hence to start to probe the developmental biology underlying the evolution of our species.

As we wrote in an editorial in January, “The human development field represents an essential growth area for the developmental biology community, and Development is keen to play an active role in supporting and inspiring it” ( Pourquié et al., 2015 ). This Special Issue celebrates that aim – bringing together a collection of Reviews and Research Articles that directly address a broad range of topics in human developmental biology: from the earliest stages of human development to cellular ageing and degeneration, and from basic questions of how an organ is formed to ways in which we might translate this knowledge in the clinic. We are also supporting this initiative with a second ‘From Stem Cells to Human Development’ meeting, to be held in September 2016. More details on what should be a fantastic follow-up event can be found at http://workshops.biologists.com/from-stem-cells-to-human-development-2/ .

Studying human development is obviously a challenging endeavour, given the practical and ethical difficulties in working with human material. However, as discussed by Dianne Gerrelli and colleagues (2015) , there is a growing set of resources for researchers, including the Human Developmental Biology Resource (with which the authors are affiliated), which provides embryonic and foetal material and a range of valuable services. Maintaining and developing such resources will be essential as research on human development progresses.

Complementing work using human tissue, much of the research into human development relies on the generation and manipulation of human pluripotent stem cells (hPSCs) – either embryonic (hESCs) or induced (hiPSCs). There has, however, been much debate surrounding the pluripotent status of such hPSCs, particularly when compared with their mouse equivalents, as well as their in vivo counterparts. In their Review, Martin Pera and colleagues ( Davidson et al., 2015 ) discuss these controversies in the light of recent attempts to generate truly naïve hESCs. Kathy Niakan and co-workers are also interested in pluripotent states and human-mouse comparisons. In their Research Article ( Blakeley et al., 2015 ), they report single-cell RNA sequencing analyses of human and mouse preimplantation epiblasts, identifying important differences in the transcriptomes – and presumably therefore the development – of the early human and mouse embryo. One challenge in the field has been that functional assays for pluripotency of human cells are limited. To address this, Hiromitsu Nakauchi and colleagues ( Masaki et al., 2015 ) investigate whether generating inter-specific chimeras (using mouse epiblasts and PSCs from various species) might provide an alternative assay system. Also using mouse embryology to probe human development are Felipe Vilella and colleagues, who describe a microRNA secreted in human endometrial fluid that can promote mouse embryo adhesion during implantation ( Vilella et al., 2015 ), potentially identifying a novel route by which the efficiency of implantation can be modulated.

In another research paper investigating the role of microRNAs in human development ( Jönsson et al., 2015 ), Malin Parmar and co-workers analyse the microRNAs expressed in the human foetal brain and in PSC-derived neural progenitor cells, identifying region-specific microRNAs that probably influence neural cell fate. Generating a functional nervous system requires not only that cell fate is correctly defined, but also that appropriate connectivity is established and that neurons are properly supported by glia. Frederick Livesey and colleagues address the former problem in cortical neuron cultures ( Kirwan et al., 2015 ), while Motoharu Sakaue and Maya Sieber-Blum describe a protocol for generating supporting Schwann-like cells from human epidermal neural crest stem cells ( Sakaue and Sieber-Blum, 2015 ). Meanwhile, Ikuo Suzuki and Pierre Vanderhaeghen (2015) review various aspects of studying neural development using hPSCs and discuss how these approaches should allow us to gain insights into the evolution of the human brain.

Katie Pollard and Lucia Franchini's interests also lie in understanding human evolution, but from a genomic perspective. Their Review ( Franchini and Pollard, 2015 ) discusses how we can combine sequencing information with functional genomics and stem cell biology to identify and characterise changes in the human genome that might have led to human-specific developmental traits. They highlight the importance of appropriate experimental systems – not only model organisms but also through human stem cells and organoids – in which to test the function of human-specific genomic features. The ability to model not only cellular differentiation but also tissue formation in a dish constitutes a major breakthrough in the field over the past decade. Meritxell Huch and Bon-Kyoung Koo review the latest advances in generating endodermal organoids from both embryonic and adult stem cells ( Huch and Koo, 2015 ) and provide a perspective on where this field is heading. The Review by Neil Hanley and colleagues ( Jennings et al., 2015 ), while also focussing on endoderm development – in this case, pancreas – provides a complementary viewpoint, discussing what is known about human pancreas development in vivo and how these insights translate into our ability to generate β-cells in vitro .

Turning to other organs, Christine Mummery and Charles Murry and their colleagues both focus on heart development. Mummery's work ( van den Berg et al., 2015 ) characterises the transcriptome of the human foetal heart and compares it with the RNA profile of PSC-derived cardiomyocytes. Meanwhile, Murry's study ( Palpant et al., 2015 ) provides insights into how cardiomyocytes are specified in a hESC system. The final Research Article of this issue returns to the topic of organoid formation, this time the mammary gland. Christina Scheel and co-workers ( Linnemann et al., 2015 ) present an organoid system that allows the regenerative potential and morphogenetic dynamics of mammary epithelial cells to be studied.

Although understanding human development is an important goal in itself, the translational potential of this field is clear: if we can grow human tissues in vitro , we can use these to model disease, to test potential drugs and to develop cell therapies. Two Spotlights in this issue discuss these aspects of the field. Scott Thies and Charles Murry (2015) present some of the most promising preclinical data and clinical trials of stem cell therapies, while Elsa Vera and Lorenz Studer (2015) highlight a potential problem with using stem cell-derived models in disease research: both hESCs and hiPSCs are ‘young’ cells, whereas many diseases – particularly neurodegenerative disorders – afflict the old. Although these articles stray from the classic scope of a developmental biology journal, we hope that they illustrate the continuum of both the field, from basic understanding of developmental processes to their applications in regenerative therapy, and of development itself – from embryogenesis through post-embryonic maturation to ageing and decline.

We have a limited number of print copies of this Special Issue to give away to interested readers. If you would like one, please send an email to [email protected] with your mailing address. Whether in print or online, we hope you enjoy this Special Issue on Human Development. We see an exciting future for this field, and we want Development to be at the heart of it. We therefore encourage those of you working in this area to consider Development as a potential venue for the publication of your best work and we look forward to many more exciting human development papers finding their way into the pages of our journal.

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A new future of work: The race to deploy AI and raise skills in Europe and beyond

At a glance.

Amid tightening labor markets and a slowdown in productivity growth, Europe and the United States face shifts in labor demand, spurred by AI and automation. Our updated modeling of the future of work finds that demand for workers in STEM-related, healthcare, and other high-skill professions would rise, while demand for occupations such as office workers, production workers, and customer service representatives would decline. By 2030, in a midpoint adoption scenario, up to 30 percent of current hours worked could be automated, accelerated by generative AI (gen AI). Efforts to achieve net-zero emissions, an aging workforce, and growth in e-commerce, as well as infrastructure and technology spending and overall economic growth, could also shift employment demand.

By 2030, Europe could require up to 12 million occupational transitions, double the prepandemic pace. In the United States, required transitions could reach almost 12 million, in line with the prepandemic norm. Both regions navigated even higher levels of labor market shifts at the height of the COVID-19 period, suggesting that they can handle this scale of future job transitions. The pace of occupational change is broadly similar among countries in Europe, although the specific mix reflects their economic variations.

Businesses will need a major skills upgrade. Demand for technological and social and emotional skills could rise as demand for physical and manual and higher cognitive skills stabilizes. Surveyed executives in Europe and the United States expressed a need not only for advanced IT and data analytics but also for critical thinking, creativity, and teaching and training—skills they report as currently being in short supply. Companies plan to focus on retraining workers, more than hiring or subcontracting, to meet skill needs.

Workers with lower wages face challenges of redeployment as demand reweights toward occupations with higher wages in both Europe and the United States. Occupations with lower wages are likely to see reductions in demand, and workers will need to acquire new skills to transition to better-paying work. If that doesn’t happen, there is a risk of a more polarized labor market, with more higher-wage jobs than workers and too many workers for existing lower-wage jobs.

Choices made today could revive productivity growth while creating better societal outcomes. Embracing the path of accelerated technology adoption with proactive worker redeployment could help Europe achieve an annual productivity growth rate of up to 3 percent through 2030. However, slow adoption would limit that to 0.3 percent, closer to today’s level of productivity growth in Western Europe. Slow worker redeployment would leave millions unable to participate productively in the future of work.

Demand will change for a range of occupations through 2030, including growth in STEM- and healthcare-related occupations, among others

This report focuses on labor markets in nine major economies in the European Union along with the United Kingdom, in comparison with the United States. Technology, including most recently the rise of gen AI, along with other factors, will spur changes in the pattern of labor demand through 2030. Our study, which uses an updated version of the McKinsey Global Institute future of work model, seeks to quantify the occupational transitions that will be required and the changing nature of demand for different types of jobs and skills.

Our methodology

We used methodology consistent with other McKinsey Global Institute reports on the future of work to model trends of job changes at the level of occupations, activities, and skills. For this report, we focused our analysis on the 2022–30 period.

Our model estimates net changes in employment demand by sector and occupation; we also estimate occupational transitions, or the net number of workers that need to change in each type of occupation, based on which occupations face declining demand by 2030 relative to current employment in 2022. We included ten countries in Europe: nine EU members—the Czech Republic, Denmark, France, Germany, Italy, Netherlands, Poland, Spain, and Sweden—and the United Kingdom. For the United States, we build on estimates published in our 2023 report Generative AI and the future of work in America.

We included multiple drivers in our modeling: automation potential, net-zero transition, e-commerce growth, remote work adoption, increases in income, aging populations, technology investments, and infrastructure investments.

Two scenarios are used to bookend the work-automation model: “late” and “early.” For Europe, we modeled a “faster” scenario and a “slower” one. For the faster scenario, we use the midpoint—the arithmetical average between our late and early scenarios. For the slower scenario, we use a “mid late” trajectory, an arithmetical average between a late adoption scenario and the midpoint scenario. For the United States, we use the midpoint scenario, based on our earlier research.

We also estimate the productivity effects of automation, using GDP per full-time-equivalent (FTE) employee as the measure of productivity. We assumed that workers displaced by automation rejoin the workforce at 2022 productivity levels, net of automation, and in line with the expected 2030 occupational mix.

Amid tightening labor markets and a slowdown in productivity growth, Europe and the United States face shifts in labor demand, spurred not only by AI and automation but also by other trends, including efforts to achieve net-zero emissions, an aging population, infrastructure spending, technology investments, and growth in e-commerce, among others (see sidebar, “Our methodology”).

Our analysis finds that demand for occupations such as health professionals and other STEM-related professionals would grow by 17 to 30 percent between 2022 and 2030, (Exhibit 1).

By contrast, demand for workers in food services, production work, customer services, sales, and office support—all of which declined over the 2012–22 period—would continue to decline until 2030. These jobs involve a high share of repetitive tasks, data collection, and elementary data processing—all activities that automated systems can handle efficiently.

Up to 30 percent of hours worked could be automated by 2030, boosted by gen AI, leading to millions of required occupational transitions

By 2030, our analysis finds that about 27 percent of current hours worked in Europe and 30 percent of hours worked in the United States could be automated, accelerated by gen AI. Our model suggests that roughly 20 percent of hours worked could still be automated even without gen AI, implying a significant acceleration.

These trends will play out in labor markets in the form of workers needing to change occupations. By 2030, under the faster adoption scenario we modeled, Europe could require up to 12.0 million occupational transitions, affecting 6.5 percent of current employment. That is double the prepandemic pace (Exhibit 2). Under a slower scenario we modeled for Europe, the number of occupational transitions needed would amount to 8.5 million, affecting 4.6 percent of current employment. In the United States, required transitions could reach almost 12.0 million, affecting 7.5 percent of current employment. Unlike Europe, this magnitude of transitions is broadly in line with the prepandemic norm.

Both regions navigated even higher levels of labor market shifts at the height of the COVID-19 period. While these were abrupt and painful to many, given the forced nature of the shifts, the experience suggests that both regions have the ability to handle this scale of future job transitions.

Businesses will need a major skills upgrade

The occupational transitions noted above herald substantial shifts in workforce skills in a future in which automation and AI are integrated into the workplace (Exhibit 3). Workers use multiple skills to perform a given task, but for the purposes of our quantification, we identified the predominant skill used.

Demand for technological skills could see substantial growth in Europe and in the United States (increases of 25 percent and 29 percent, respectively, in hours worked by 2030 compared to 2022) under our midpoint scenario of automation adoption (which is the faster scenario for Europe).

Demand for social and emotional skills could rise by 11 percent in Europe and by 14 percent in the United States. Underlying this increase is higher demand for roles requiring interpersonal empathy and leadership skills. These skills are crucial in healthcare and managerial roles in an evolving economy that demands greater adaptability and flexibility.

Conversely, demand for work in which basic cognitive skills predominate is expected to decline by 14 percent. Basic cognitive skills are required primarily in office support or customer service roles, which are highly susceptible to being automated by AI. Among work characterized by these basic cognitive skills experiencing significant drops in demand are basic data processing and literacy, numeracy, and communication.

Demand for work in which higher cognitive skills predominate could also decline slightly, according to our analysis. While creativity is expected to remain highly sought after, with a potential increase of 12 percent by 2030, work activities characterized by other advanced cognitive skills such as advanced literacy and writing, along with quantitative and statistical skills, could decline by 19 percent.

Demand for physical and manual skills, on the other hand, could remain roughly level with the present. These skills remain the largest share of workforce skills, representing about 30 percent of total hours worked in 2022. Growth in demand for these skills between 2022 and 2030 could come from the build-out of infrastructure and higher investment in low-emissions sectors, while declines would be in line with continued automation in production work.

Business executives report skills shortages today and expect them to worsen

A survey we conducted of C-suite executives in five countries shows that companies are already grappling with skills challenges, including a skills mismatch, particularly in technological, higher cognitive, and social and emotional skills: about one-third of the more than 1,100 respondents report a shortfall in these critical areas. At the same time, a notable number of executives say they have enough employees with basic cognitive skills and, to a lesser extent, physical and manual skills.

Within technological skills, companies in our survey reported that their most significant shortages are in advanced IT skills and programming, advanced data analysis, and mathematical skills. Among higher cognitive skills, significant shortfalls are seen in critical thinking and problem structuring and in complex information processing. About 40 percent of the executives surveyed pointed to a shortage of workers with these skills, which are needed for working alongside new technologies (Exhibit 4).

Companies see retraining as key to acquiring needed skills and adapting to the new work landscape

Surveyed executives expect significant changes to their workforce skill levels and worry about not finding the right skills by 2030. More than one in four survey respondents said that failing to capture the needed skills could directly harm financial performance and indirectly impede their efforts to leverage the value from AI.

To acquire the skills they need, companies have three main options: retraining, hiring, and contracting workers. Our survey suggests that executives are looking at all three options, with retraining the most widely reported tactic planned to address the skills mismatch: on average, out of companies that mentioned retraining as one of their tactics to address skills mismatch, executives said they would retrain 32 percent of their workforce. The scale of retraining needs varies in degree. For example, respondents in the automotive industry expect 36 percent of their workforce to be retrained, compared with 28 percent in the financial services industry. Out of those who have mentioned hiring or contracting as their tactics to address the skills mismatch, executives surveyed said they would hire an average of 23 percent of their workforce and contract an average of 18 percent.

Occupational transitions will affect high-, medium-, and low-wage workers differently

All ten European countries we examined for this report may see increasing demand for top-earning occupations. By contrast, workers in the two lowest-wage-bracket occupations could be three to five times more likely to have to change occupations compared to the top wage earners, our analysis finds. The disparity is much higher in the United States, where workers in the two lowest-wage-bracket occupations are up to 14 times more likely to face occupational shifts than the highest earners. In Europe, the middle-wage population could be twice as affected by occupational transitions as the same population in United States, representing 7.3 percent of the working population who might face occupational transitions.

Enhancing human capital at the same time as deploying the technology rapidly could boost annual productivity growth

About quantumblack, ai by mckinsey.

QuantumBlack, McKinsey’s AI arm, helps companies transform using the power of technology, technical expertise, and industry experts. With thousands of practitioners at QuantumBlack (data engineers, data scientists, product managers, designers, and software engineers) and McKinsey (industry and domain experts), we are working to solve the world’s most important AI challenges. QuantumBlack Labs is our center of technology development and client innovation, which has been driving cutting-edge advancements and developments in AI through locations across the globe.

Organizations and policy makers have choices to make; the way they approach AI and automation, along with human capital augmentation, will affect economic and societal outcomes.

We have attempted to quantify at a high level the potential effects of different stances to AI deployment on productivity in Europe. Our analysis considers two dimensions. The first is the adoption rate of AI and automation technologies. We consider the faster scenario and the late scenario for technology adoption. Faster adoption would unlock greater productivity growth potential but also, potentially, more short-term labor disruption than the late scenario.

The second dimension we consider is the level of automated worker time that is redeployed into the economy. This represents the ability to redeploy the time gained by automation and productivity gains (for example, new tasks and job creation). This could vary depending on the success of worker training programs and strategies to match demand and supply in labor markets.

We based our analysis on two potential scenarios: either all displaced workers would be able to fully rejoin the economy at a similar productivity level as in 2022 or only some 80 percent of the automated workers’ time will be redeployed into the economy.

Exhibit 5 illustrates the various outcomes in terms of annual productivity growth rate. The top-right quadrant illustrates the highest economy-wide productivity, with an annual productivity growth rate of up to 3.1 percent. It requires fast adoption of technologies as well as full redeployment of displaced workers. The top-left quadrant also demonstrates technology adoption on a fast trajectory and shows a relatively high productivity growth rate (up to 2.5 percent). However, about 6.0 percent of total hours worked (equivalent to 10.2 million people not working) would not be redeployed in the economy. Finally, the two bottom quadrants depict the failure to adopt AI and automation, leading to limited productivity gains and translating into limited labor market disruptions.

Four priorities for companies

The adoption of automation technologies will be decisive in protecting businesses’ competitive advantage in an automation and AI era. To ensure successful deployment at a company level, business leaders can embrace four priorities.

Understand the potential. Leaders need to understand the potential of these technologies, notably including how AI and gen AI can augment and automate work. This includes estimating both the total capacity that these technologies could free up and their impact on role composition and skills requirements. Understanding this allows business leaders to frame their end-to-end strategy and adoption goals with regard to these technologies.

Plan a strategic workforce shift. Once they understand the potential of automation technologies, leaders need to plan the company’s shift toward readiness for the automation and AI era. This requires sizing the workforce and skill needs, based on strategically identified use cases, to assess the potential future talent gap. From this analysis will flow details about the extent of recruitment of new talent, upskilling, or reskilling of the current workforce that is needed, as well as where to redeploy freed capacity to more value-added tasks.

Prioritize people development. To ensure that the right talent is on hand to sustain the company strategy during all transformation phases, leaders could consider strengthening their capabilities to identify, attract, and recruit future AI and gen AI leaders in a tight market. They will also likely need to accelerate the building of AI and gen AI capabilities in the workforce. Nontechnical talent will also need training to adapt to the changing skills environment. Finally, leaders could deploy an HR strategy and operating model to fit the post–gen AI workforce.

Pursue the executive-education journey on automation technologies. Leaders also need to undertake their own education journey on automation technologies to maximize their contributions to their companies during the coming transformation. This includes empowering senior managers to explore automation technologies implications and subsequently role model to others, as well as bringing all company leaders together to create a dedicated road map to drive business and employee value.

AI and the toolbox of advanced new technologies are evolving at a breathtaking pace. For companies and policy makers, these technologies are highly compelling because they promise a range of benefits, including higher productivity, which could lift growth and prosperity. Yet, as this report has sought to illustrate, making full use of the advantages on offer will also require paying attention to the critical element of human capital. In the best-case scenario, workers’ skills will develop and adapt to new technological challenges. Achieving this goal in our new technological age will be highly challenging—but the benefits will be great.

Eric Hazan is a McKinsey senior partner based in Paris; Anu Madgavkar and Michael Chui are McKinsey Global Institute partners based in New Jersey and San Francisco, respectively; Sven Smit is chair of the McKinsey Global Institute and a McKinsey senior partner based in Amsterdam; Dana Maor is a McKinsey senior partner based in Tel Aviv; Gurneet Singh Dandona is an associate partner and a senior expert based in New York; and Roland Huyghues-Despointes is a consultant based in Paris.

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Recycling of printed Xerographic paper using Aspergillus assiutensis enzyme cocktail: an integrated approach to sustainable development

• Research Article
• Published: 30 May 2024

Cite this article

• Aiman Tanveer 1   na1 ,
• Supriya Gupta 1   na1 ,
• Shruti Dwivedi 1 ,
• Sangeeta Yadav 1 &
• Dinesh Yadav   ORCID: orcid.org/0000-0003-2990-0670 1  

To overcome the human and animal survivability risk, sustainable development is the only option on earth that can be achieved through the maximum use of renewable environmental resources. Recycling of waste paper is an emerging waste management approach to conserve natural resources. Herein, we studied enzyme-mediated process to recycle the xerographic paper by using the crude fungal extract from indigenously isolated fungi identified as  Aspergillus assiutensis . The fungal enzyme cocktail has been characterized for the production of multiple enzymes namely cellulase, amylase, xylanase, pectinase, and protease. All these enzymes have pH optima in the acidic range and except cellulase and all the enzymes are stable from 10 to 80 C. In the zymogram analysis, pectinase, xylanase, amylase, and cellulase were detected at 68 kDa, ~ 54 kDa, 38 kDa, and 30 kDa, respectively. Also, the presence of protease was confirmed by the clear zone at 68, 31, and 16 kDa. A 26% decrease in the kappa number and reduction in Hex A of the pulp was observed on the treatment of the pulp with enzyme as compared to the control pulp without any treatment. The physical and chemical properties of the pulp were also improved by enzyme-mediated pulping as compared to the control The physiochemical parameter of the effluent like TDS was reduced (397 ppm) significantly in comparison to chemical deinking process and it was within the permissible limit. BOD and alkalinity were reduced when the enzymes and chemical dosage were used in combination. These results indicate that chemi-enzymatic deinking is most promising to reduce or remove the pollution parameters including ink and this approach can be used in the paper and pulp industry for sustainable development.

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The datasets used and analyzed during the current study are available from the corresponding author on reasonable request.

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Acknowledgements

DST Women Scientist Scheme-B (Grant. No. DST/WOS-B/2018/1707) to Dr. Aiman Tanveer is acknowledged.

Funding provided by the Department of Science and Technology, New Delhi in the form of the DST Women Scientist Scheme-B (Grant. No. DST/WOS-B/2018/1707) to Dr. Aiman Tanveer is acknowledged.

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Department of Biotechnology, Deen Dayal Upadhyaya Gorakhpur University, Gorakhpur, 273009, Uttar Pradesh, India

Aiman Tanveer, Supriya Gupta, Shruti Dwivedi, Sangeeta Yadav & Dinesh Yadav

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AT prepared the overall first draft and conceptualized it. SG performed the experiments and compiled the results. SD prepared the Figure and Tables. SY reviewed and corrected the first draft of the manuscript. DY supervised the research work, finalized the overall content, improved the language, and prepared the final draft for submission.

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Tanveer, A., Gupta, S., Dwivedi, S. et al. Recycling of printed Xerographic paper using Aspergillus assiutensis enzyme cocktail: an integrated approach to sustainable development. Environ Sci Pollut Res (2024). https://doi.org/10.1007/s11356-024-33780-2

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Received : 26 August 2023

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Published : 30 May 2024

DOI : https://doi.org/10.1007/s11356-024-33780-2

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