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The identification of distinct waves of progenitors during development, each corresponding to a specific time, space, and function, provided the basis for the concept of a “layered” organization in development. The concept of a layered hematopoiesis was established by classical embryology studies in birds and amphibians. Recent progress in generating reliable lineage tracing models together with transcriptional and proteomic analyses in single cells revealed that, also in mammals, the hematopoietic system evolves in successive waves of progenitors with distinct properties and fate. During embryogenesis, sequential waves of hematopoietic progenitors emerge at different anatomic sites, generating specific cell types with distinct functions and tissue homing capacities. The first progenitors originate in the yolk sac before the emergence of hematopoietic stem cells, some giving rise to progenies that persist throughout life. Hematopoietic stem cell-derived cells that protect organisms against environmental pathogens follow the same sequential strategy, with subsets of lymphoid cells being only produced during embryonic development. Growing evidence indicates that fetal immune cells contribute to the proper development of the organs they seed and later ensure life-long tissue homeostasis and immune protection. They include macrophages, mast cells, some γδ T cells, B-1 B cells, and innate lymphoid cells, which have “non-redundant” functions, and early perturbations in their development or function affect immunity in the adult. These observations challenged the view that all hematopoietic cells found in the adult result from constant and monotonous production from bone marrow-resident hematopoietic stem cells. In this review, we evaluate evidence for a layered hematopoietic system across species. We discuss mechanisms and selective pressures leading to the temporal generation of different cell types. We elaborate on the consequences of disturbing fetal immune cells on tissue homeostasis and immune development later in life.

Abstract: The emerging science of evolutionary developmental (“evo devo”) biology can aid us in thinking about our universe as both an evolutionary system, where most processes are unpredictable and creative, and a developmental system, where a special few processes are predictable and constrained to produce far-future-specific emergent order, just as we see in the common developmental processes in two stars of an identical population type, or in two genetically identical twins in biology. The transcension hypothesis proposes that a universal process of evolutionary development guides all sufficiently advanced civilizations into what may be called "inner space," a computationally optimal domain of increasingly dense, productive, miniaturized, and efficient scales of space, time, energy, and matter, and eventually, to a black-hole-like destination. Transcension as a developmental destiny might also contribute to the solution to the Fermi paradox, the question of why we have not seen evidence of or received beacons from intelligent civilizations. A few potential evolutionary, developmental, and information theoretic reasons, mechanisms, and models for constrained transcension of advanced intelligence are briefly considered. In particular, we introduce arguments that black holes may be a developmental destiny and standard attractor for all higher intelligence, as they appear to some to be ideal computing, learning, forward time travel, energy harvesting, civilization merger, natural selection, and universe replication devices. In the transcension hypothesis, simpler civilizations that succeed in resisting transcension by staying in outer (normal) space would be developmental failures, which are statistically very rare late in the life cycle of any biological developing system. If transcension is a developmental process, we may expect brief broadcasts or subtle forms of galactic engineering to occur in small portions of a few galaxies, the handiwork of young and immature civilizations, but constrained transcension should be by far the norm for all mature civilizations. The transcension hypothesis has significant and testable implications for our current and future METI and SETI agendas. If all universal intelligence eventually transcends to black-hole-like environments, after which some form of merger and selection occurs, and if two-way messaging (a send–receive cycle) is severely limited by the great distances between neighboring and rapidly transcending civilizations, then sending one-way METI or probes prior to transcension becomes the only real communication option. But one-way messaging or probes may provably reduce the evolutionary diversity in all civilizations receiving the message, as they would then arrive at their local transcensions in a much more homogenous fashion. If true, an ethical injunction against one-way messaging or probes might emerge in the morality and sustainability systems of all sufficiently advanced civilizations, an argument known as the Zoo hypothesis in Fermi paradox literature, if all higher intelligences are subject to an evolutionary attractor to maximize their local diversity, and a developmental attractor to merge and advance universal intelligence. In any such environment, the evolutionary value of sending any interstellar message or probe may simply not be worth the cost, if transcension is an inevitable, accelerative, and testable developmental process, one that eventually will be discovered and quantitatively described by future physics. Fortunately, transcension processes may be measurable today even without good physical theory, and radio and optical SETI may each provide empirical tests. If transcension is a universal developmental constraint, then without exception all early and low-power electromagnetic leakage signals (radar, radio, television), and later, optical evidence of the exoplanets and their atmospheres should reliably cease as each civilization enters its own technological singularities (emergence of postbiological intelligence and life forms) and recognizes that they are on an optimal and accelerating path to a black-hole-like environment. Furthermore, optical SETI may soon allow us to map an expanding area of the galactic habitable zone we may call the galactic transcension zone, an inner ring that contains older transcended civilizations, and a missing planets problem as we discover that planets with life signatures occur at a much lower frequencies in this inner ring than in the remainder of the habitable zone. [Copyright &y& Elsevier]

The identification of distinct waves of progenitors during development, each corresponding to a specific time, space, and function, provided the basis for the concept of a "layered" organization in development. The concept of a layered hematopoiesis was established by classical embryology studies in birds and amphibians. Recent progress in generating reliable lineage tracing models together with transcriptional and proteomic analyses in single cells revealed that, also in mammals, the hematopoietic system evolves in successive waves of progenitors with distinct properties and fate. During embryogenesis, sequential waves of hematopoietic progenitors emerge at different anatomic sites, generating specific cell types with distinct functions and tissue homing capacities. The first progenitors originate in the yolk sac before the emergence of hematopoietic stem cells, some giving rise to progenies that persist throughout life. Hematopoietic stem cell-derived cells that protect organisms against environmental pathogens follow the same sequential strategy, with subsets of lymphoid cells being only produced during embryonic development. Growing evidence indicates that fetal immune cells contribute to the proper development of the organs they seed and later ensure life-long tissue homeostasis and immune protection. They include macrophages, mast cells, some γδ T cells, B-1 B cells, and innate lymphoid cells, which have "non-redundant" functions, and early perturbations in their development or function affect immunity in the adult. These observations challenged the view that all hematopoietic cells found in the adult result from constant and monotonous production from bone marrow-resident hematopoietic stem cells. In this review, we evaluate evidence for a layered hematopoietic system across species. We discuss mechanisms and selective pressures leading to the temporal generation of different cell types. We elaborate on the consequences of disturbing fetal immune cells on tissue homeostasis and immune development later in life., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2020 Elsaid, Soares-da-Silva, Peixoto, Amiri, Mackowski, Pereira, Bandeira and Cumano.)