ArticlePDF AvailableLiterature Review

Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos

Authors:
Edward J Steele at CY O'Connor ERADE Village Foundation
Edward J Steele
  • CY O'Connor ERADE Village Foundation
Reginald Gorczynski at University of Toronto
Reginald Gorczynski
Robyn A. Lindley
Robyn A. Lindley
Robyn A. Lindley
  • This person is not on ResearchGate, or hasn't claimed this research yet.
Yongsheng Liu
Yongsheng Liu
Yongsheng Liu
  • This person is not on ResearchGate, or hasn't claimed this research yet.
Show all 8 authorsHide
Download full-text PDF
Read full-text
Download citation

Abstract and Figures

We review the main lines of evidence (molecular, cellular and whole organism) published since the 1970s demonstrating Lamarckian Inheritance in animals, plants and microorganisms viz. the transgenerational inheritance of environmentally-induced acquired characteristics. The studies in animals demonstrate the genetic permeability of the soma-germline Weismann Barrier. The widespread nature of environmentally-directed inheritance phenomena reviewed here contradicts a key pillar of neo-Darwinism which affirms the rigidity of the Weismann Barrier. These developments suggest that neo-Darwinian evolutionary theory is in need of significant revision. We argue that Lamarckian inheritance strategies involving environmentally-induced rapid directional genetic adaptations make biological sense in the context of cosmic Panspermia allowing the efficient spread of living systems and genetic innovation throughout the Universe. The Hoyle-Wickramasinghe Panspermia paradigm also developed since the 1970s, unlike strictly geocentric neo-Darwinism provides a cogent biological rationale for the actual widespread existence of Lamarckian modes of inheritance - it provides its raison d'être. Under a terrestrially confined neo-Darwinian viewpoint such an association may have been thought spurious in the past. Our aim is to outline the conceptual links between rapid Lamarckian-based evolutionary hypermutation processes dependent on reverse transcription-coupled mechanisms among others and the effective cosmic spread of living systems. For example, a viable, or cryo-preserved, living system travelling through space in a protective matrix will need of necessity to rapidly adapt and proliferate on landing in a new cosmic niche. Lamarckian mechanisms thus come to the fore and supersede the slow (blind and random) genetic processes expected under a traditional neo-Darwinian evolutionary paradigm.
The key features of the reverse transcriptase mechanism of somatic hypermutation (SHM) at Transcription Bubbles -a type of representation of the deaminase-based "Universal Mutator" likely to operate in many kingdoms of life (Lindley, 2018; Krishnan et al., 2018). Some elements of this figure have appeared before, and this figure is a modified combination of parts from (Steele, 2017), Lindley and Steele (2013), as well as from mechanism figures in Steele (2009, 2016a) and Steele and Lindley (2017). This is also an adaptation of the target site reverse transcription process reported in Luan et al. (1993). Shown is an RNA Polymerase II-generated Transcription Bubble with C-site and A-site substrate deamination events by AID, APOBEC and ADAR deaminase enzymes, which generates the strand-biased transition mutation signatures -A-to-G, G-to-A, G-to-T, and G-to-C. DNA strands shown by black lines; pre-mRNA as red lines; cDNA strands as thick blue lines due to DNA polymerase
The key features of the reverse transcriptase mechanism of somatic hypermutation (SHM) at Transcription Bubbles -a type of representation of the deaminase-based "Universal Mutator" likely to operate in many kingdoms of life (Lindley, 2018; Krishnan et al., 2018). Some elements of this figure have appeared before, and this figure is a modified combination of parts from (Steele, 2017), Lindley and Steele (2013), as well as from mechanism figures in Steele (2009, 2016a) and Steele and Lindley (2017). This is also an adaptation of the target site reverse transcription process reported in Luan et al. (1993). Shown is an RNA Polymerase II-generated Transcription Bubble with C-site and A-site substrate deamination events by AID, APOBEC and ADAR deaminase enzymes, which generates the strand-biased transition mutation signatures -A-to-G, G-to-A, G-to-T, and G-to-C. DNA strands shown by black lines; pre-mRNA as red lines; cDNA strands as thick blue lines due to DNA polymerase
… 
Coupled Epigenetic-Genetic Mechanisms of Lamarckian Inheritance with the Penetration of the Weismann Barrier in Higher Animals. The point of the sequence is to show that genes targeted first for epigenetic transgenerational regulation can mature via a reverse transcription step targeting the same genes causing hard genetic (DNA) inheritance. The key epigenetic-genetic concepts are discussed at length in Lindley (2010, 2011) and specifically in Lindley (2018). The properties of cytosine modifications play a key role in the plasticity of the Epigenetic-Genetic coupling (see Guo et al., 2011a,b, Nabel et al., 2012). The distinction between "soft" and "hard" Lamarckian inheritance is discussed explicitly in Steele (2016b). The often confused distinction between the Central Dogma of Molecular Biology and the Weismann Barrier (Steele, 1979; 2016b) is discussed and further clarified in Noble (2018). In the first phase after stimulation, the environmentally-induced "epigenetic" gene regulatory factors (e.g. methylation-demethylation at CpG sites and other modifications at cytosines; synthesis of small 21 nt-24nt sRNAs such as miRNAs etc. and guided by other non-coding regulatory RNAs) light up and target specific genes and gene pathways for regulated gene expression. The epigenetic role of long non-coding RNAs (>200 nt) targeting regulatory portions flanking protein-coding genes is covered in conceptual detail by Mattick (2003, 2018). The recent review by Kulski (2019) shows the functional importance of locus-wide lncRNAs in conserving long ancestral haplotypes at the human Major Histocompatibility Complex locus -where they act as genomic anchor points for binding transcription factors, enhancers, and chromatin remodeling enzymes thus regulating transcription and chromatin folding. LncRNAs specifically target DNA sequences usually via RNA-DNA triple helix interactions involving the weaker yet biologically significant Hoogsteen hydrogen bonding. Hoogsteen base pairing, considerably weaker than Watson-Crick base pairing is varied, in both parallel and anti-parallel configurations with the RNA sequence aligned in the major groove of the DNA duplex (Li et al., 2016). These allow multiple points of hydrogen bonding over significant sequence lengths (e.g. Enhancer or Promoter regions) thus allowing gene-specific recognition. RNA-DNA triple helix interactions thus allow targeted delivery of chromatin modifications resulting in either active transcription (activation via acetyltransferase-associated complexes) or gene silencing (chromatin compaction via methyltransferase -associated complexes). These data and the analytical methodology are reviewed in detail in Smith et al. (2013, 2017), Buske et al., (2011, 2012) and in Li et al. (2016). Such lncRNA epigenetic regulators are ubiquitously found in secreted extracellular vesicles and exosomes, particularly in tumours and tumour cell microenvironments (Xie et al., 2019; Chen et al., 2019). Epigenetically "marked" genes can become targets for AID/APOBEC-deaminase mediated cytosine to uracil (C-to-U) and cytosine to thymine (C-to-T) mutations (at 5Me CpG sites, Morgan et al., 2004), which result in GU and GT mispairs. Then as shown in part in Fig. 1 these can progress through further error-prone steps of DNA repair following base excision resulting in Abasic sites and then single stranded DNA nicks in the transcribed strand with 3 0 OH ends that can prime both DNA and RNA-dependent cDNA synthesis (off homologous newly transcribed RNA sequence templates). These downstream nicks resulting from ncRNA regulatory targeting therefore "open" the DNA in that genomic region to invasion and targeting of previously base-modified and mutated mRNA sequence templates which can be reverse transcribed and their specific cDNA fragments integrated at these C-sites (and surrounding sequence) into the genomic DNA, by target site reverse transcription,TSRT (Luan et al., 1993) as discussed at length elsewhere (e.g. Steele, 2016a. Steele and Lindley, 2017) and shown in Fig. 1. This is a variant of AID/APOBEC deaminase-mediated dysregulated immunoglobulin somatic hypermutation-like responses initiated at C-sites across the cancer genome (Lindley, 2013; Lindley and Steele, 2013; Lindley et al., 2016). ADAR deaminases causing adenosine to inosine modifications in RNA and DNA are also part of this dysregulated Ig SHM-like response scheme (A-to-I, read out as A-to-G transitions, Lindley, 2013; Lindley and Steele, 2013; Steele and Lindley, 2017). The enzymatic deaminase targeting specific C-sites and Asites in DNA and RNA substrates occurs in protein-coding regions in codon context (Lindley, 2013) most plausibly in the 3D environment of stalled Transcription Bubbles (Lindley, 2013; Steele and Lindley, 2017). There is evidence that deamination of 5-methylcytosine (5 mC) and 5-hydroxymethylcytosine (5hmC) and generation of mutagenic C-to-T mutations directly by the activity of AID/APOBEC complexes is an alternative path to successive oxidation reactions by TET enzymes for the initiation and regulation of DNA demethylation (Guo et al., 2011a,b, Nabel et al., 2012, Pastor et al., 2013, Scourzic et al., 2015). The role of extracellular secreted vesicles and exosomes is discussed in Fig. 1 of Steele and Lloyd (2015) based on the seminal vesicle/exosome data published in Cossetti et al. (2014) (mice) and later in Sharma et al. (2016) (humans). B lymphocytes themselves when activated by mitogens secrete large numbers of endogenous retroviruses (Moroni and Schumann, 1975; Moroni et al., 1980). The significance of the very high concentrations (>10 11 per ml) of endogenous retroviruses in seminal fluid, surrounding the placenta and actually bound to the heads of spermatozoa (Keissling et al., 1987) challenges the widely held belief that only one successful sperm affects the internalized genetic cargo at fertilization. The questioning of this commonly held belief is justified given the huge number of spermatozoa attached to a given ovum. Finally, apoptotic vesicles have also been invoked as DNA/RNA soma-to-germline transmission vehicles (Steele et al., 2002). Indeed there are many formal similarities between retroviruses and secreted extracellular vesicles (Hoena et al., 2016). The properties of extracellular extruded vesicles and exosomes has been extensively reviewed (van der Pol et al., 2012) and extracellular membrane vesicles with exported cargos appear across the three domains of life and form a intercellular communication system (Gill et al., 2018). All this has been recently highlighted by Noble (2019) in the context of Darwin's Pangenesis.
Coupled Epigenetic-Genetic Mechanisms of Lamarckian Inheritance with the Penetration of the Weismann Barrier in Higher Animals. The point of the sequence is to show that genes targeted first for epigenetic transgenerational regulation can mature via a reverse transcription step targeting the same genes causing hard genetic (DNA) inheritance. The key epigenetic-genetic concepts are discussed at length in Lindley (2010, 2011) and specifically in Lindley (2018). The properties of cytosine modifications play a key role in the plasticity of the Epigenetic-Genetic coupling (see Guo et al., 2011a,b, Nabel et al., 2012). The distinction between "soft" and "hard" Lamarckian inheritance is discussed explicitly in Steele (2016b). The often confused distinction between the Central Dogma of Molecular Biology and the Weismann Barrier (Steele, 1979; 2016b) is discussed and further clarified in Noble (2018). In the first phase after stimulation, the environmentally-induced "epigenetic" gene regulatory factors (e.g. methylation-demethylation at CpG sites and other modifications at cytosines; synthesis of small 21 nt-24nt sRNAs such as miRNAs etc. and guided by other non-coding regulatory RNAs) light up and target specific genes and gene pathways for regulated gene expression. The epigenetic role of long non-coding RNAs (>200 nt) targeting regulatory portions flanking protein-coding genes is covered in conceptual detail by Mattick (2003, 2018). The recent review by Kulski (2019) shows the functional importance of locus-wide lncRNAs in conserving long ancestral haplotypes at the human Major Histocompatibility Complex locus -where they act as genomic anchor points for binding transcription factors, enhancers, and chromatin remodeling enzymes thus regulating transcription and chromatin folding. LncRNAs specifically target DNA sequences usually via RNA-DNA triple helix interactions involving the weaker yet biologically significant Hoogsteen hydrogen bonding. Hoogsteen base pairing, considerably weaker than Watson-Crick base pairing is varied, in both parallel and anti-parallel configurations with the RNA sequence aligned in the major groove of the DNA duplex (Li et al., 2016). These allow multiple points of hydrogen bonding over significant sequence lengths (e.g. Enhancer or Promoter regions) thus allowing gene-specific recognition. RNA-DNA triple helix interactions thus allow targeted delivery of chromatin modifications resulting in either active transcription (activation via acetyltransferase-associated complexes) or gene silencing (chromatin compaction via methyltransferase -associated complexes). These data and the analytical methodology are reviewed in detail in Smith et al. (2013, 2017), Buske et al., (2011, 2012) and in Li et al. (2016). Such lncRNA epigenetic regulators are ubiquitously found in secreted extracellular vesicles and exosomes, particularly in tumours and tumour cell microenvironments (Xie et al., 2019; Chen et al., 2019). Epigenetically "marked" genes can become targets for AID/APOBEC-deaminase mediated cytosine to uracil (C-to-U) and cytosine to thymine (C-to-T) mutations (at 5Me CpG sites, Morgan et al., 2004), which result in GU and GT mispairs. Then as shown in part in Fig. 1 these can progress through further error-prone steps of DNA repair following base excision resulting in Abasic sites and then single stranded DNA nicks in the transcribed strand with 3 0 OH ends that can prime both DNA and RNA-dependent cDNA synthesis (off homologous newly transcribed RNA sequence templates). These downstream nicks resulting from ncRNA regulatory targeting therefore "open" the DNA in that genomic region to invasion and targeting of previously base-modified and mutated mRNA sequence templates which can be reverse transcribed and their specific cDNA fragments integrated at these C-sites (and surrounding sequence) into the genomic DNA, by target site reverse transcription,TSRT (Luan et al., 1993) as discussed at length elsewhere (e.g. Steele, 2016a. Steele and Lindley, 2017) and shown in Fig. 1. This is a variant of AID/APOBEC deaminase-mediated dysregulated immunoglobulin somatic hypermutation-like responses initiated at C-sites across the cancer genome (Lindley, 2013; Lindley and Steele, 2013; Lindley et al., 2016). ADAR deaminases causing adenosine to inosine modifications in RNA and DNA are also part of this dysregulated Ig SHM-like response scheme (A-to-I, read out as A-to-G transitions, Lindley, 2013; Lindley and Steele, 2013; Steele and Lindley, 2017). The enzymatic deaminase targeting specific C-sites and Asites in DNA and RNA substrates occurs in protein-coding regions in codon context (Lindley, 2013) most plausibly in the 3D environment of stalled Transcription Bubbles (Lindley, 2013; Steele and Lindley, 2017). There is evidence that deamination of 5-methylcytosine (5 mC) and 5-hydroxymethylcytosine (5hmC) and generation of mutagenic C-to-T mutations directly by the activity of AID/APOBEC complexes is an alternative path to successive oxidation reactions by TET enzymes for the initiation and regulation of DNA demethylation (Guo et al., 2011a,b, Nabel et al., 2012, Pastor et al., 2013, Scourzic et al., 2015). The role of extracellular secreted vesicles and exosomes is discussed in Fig. 1 of Steele and Lloyd (2015) based on the seminal vesicle/exosome data published in Cossetti et al. (2014) (mice) and later in Sharma et al. (2016) (humans). B lymphocytes themselves when activated by mitogens secrete large numbers of endogenous retroviruses (Moroni and Schumann, 1975; Moroni et al., 1980). The significance of the very high concentrations (>10 11 per ml) of endogenous retroviruses in seminal fluid, surrounding the placenta and actually bound to the heads of spermatozoa (Keissling et al., 1987) challenges the widely held belief that only one successful sperm affects the internalized genetic cargo at fertilization. The questioning of this commonly held belief is justified given the huge number of spermatozoa attached to a given ovum. Finally, apoptotic vesicles have also been invoked as DNA/RNA soma-to-germline transmission vehicles (Steele et al., 2002). Indeed there are many formal similarities between retroviruses and secreted extracellular vesicles (Hoena et al., 2016). The properties of extracellular extruded vesicles and exosomes has been extensively reviewed (van der Pol et al., 2012) and extracellular membrane vesicles with exported cargos appear across the three domains of life and form a intercellular communication system (Gill et al., 2018). All this has been recently highlighted by Noble (2019) in the context of Darwin's Pangenesis.
… 
Graft-induced variations in eggplant, in which 9-leaf eggplant was grafted onto white eggplant. First row: left: fruit of white eggplant; Right: fruit of 9-leaf eggplant; Middle: fruit in the stock of the immediate generation. Second row: Variant fruit in F1 generation. Third-Fifth row: Variant fruits in the F2 generation. Reproduced with permission from Zu, D.-M., Zhao, Y.-S. (1957) A study on the vegetative hybridization of some Solanaceous plants. Scientia Sinica, 6, 889e903. From Fig. 5 in Liu (2018).
Graft-induced variations in eggplant, in which 9-leaf eggplant was grafted onto white eggplant. First row: left: fruit of white eggplant; Right: fruit of 9-leaf eggplant; Middle: fruit in the stock of the immediate generation. Second row: Variant fruit in F1 generation. Third-Fifth row: Variant fruits in the F2 generation. Reproduced with permission from Zu, D.-M., Zhao, Y.-S. (1957) A study on the vegetative hybridization of some Solanaceous plants. Scientia Sinica, 6, 889e903. From Fig. 5 in Liu (2018).
… 
The surface reflectivity spectra of comet 67P/C-G (left panel, Capaccione et al., 2015; taken at different times) compared with the transmittance spectrum of desiccated E-coli (cf. Figs. 5 and 6) over approximately the same wavelength range for comparison. Also from Wickramasinghe et al., 2018a,b).
The surface reflectivity spectra of comet 67P/C-G (left panel, Capaccione et al., 2015; taken at different times) compared with the transmittance spectrum of desiccated E-coli (cf. Figs. 5 and 6) over approximately the same wavelength range for comparison. Also from Wickramasinghe et al., 2018a,b).
… 
Amplification cycle of cosmic life. Within our galaxy alone about 100 billion circuits have been completed, one for every sun-like star.
+2
Amplification cycle of cosmic life. Within our galaxy alone about 100 billion circuits have been completed, one for every sun-like star.
… 
Figures - uploaded by Robert Temple
Author content
All figure content in this area was uploaded by Robert Temple
Content may be subject to copyright.
Content uploaded by Robert Temple
Author content
All content in this area was uploaded by Robert Temple on Oct 15, 2019
Content may be subject to copyright.
Lamarck and Panspermia - On the Efficient Spread of Living Systems
Throughout the Cosmos
Edward J. Steele
a
,
b
,
c
,
*
, Reginald M. Gorczynski
d
, Robyn A. Lindley
e
,
f
, Yongsheng Liu
g
,
Robert Temple
h
, Gensuke Tokoro
b
,
i
, Dayal T. Wickramasinghe
b
,
j
,
N. Chandra Wickramasinghe
b
,
i
,
k
a
C.Y.O'Connor ERADE Village Foundation, Piara Waters, Perth, 6112, WA, Australia
b
Centre for Astrobiology, University of Ruhuna, Matara, Sri Lanka
c
Melville Analytics Pty Ltd, Melbourne, Vic, Australia
d
Department of Surgery &Immunology, University of Toronto, Toronto, Ontario, Canada
e
Department of Clinical Pathology, Faculty of Medicine, Dentistry &Health Sciences, University of MelbourneVic, Australia
f
GMDx Group Ltd, Melbourne, Vic, Australia
g
Henan Collaborative Innovation Center of Modern Biological Breeding, Henan Institute of Science and Technology, Xinxiang, 453003, China
h
The History of Chinese Science and Culture Foundation, Conway Hall, London, UK
i
Institute for the Study of Panspermia and Astrobiology, Gifu, Japan
j
College of Physical and Mathematical Sciences, Australian National University, Canberra, Australia
k
Buckingham Centre for Astrobiology, University of Buckingham, UK
article info
Article history:
Received 18 May 2019
Received in revised form
14 August 2019
Accepted 21 August 2019
Available online xxx
Keywords:
Panspermia
Interstellar dust
Comets
Lamarckian inheritance
Neo-Darwinism
Epigenetic-genetic coupling
Reverse transcription-linked somatic
hypermutation
abstract
We review the main lines of evidence (molecular, cellular and whole organism) published since the
1970s demonstrating Lamarckian Inheritance in animals, plants and microorganisms viz. the trans-
generational inheritance of environmentally-induced acquired characteristics. The studies in animals
demonstrate the genetic permeability of the soma-germline Weismann Barrier. The widespread nature of
environmentally-directed inheritance phenomena reviewed here contradicts a key pillar of neo-
Darwinism which affirms the rigidity of the Weismann Barrier. These developments suggest that neo-
Darwinian evolutionary theory is in need of significant revision. We argue that Lamarckian inheri-
tance strategies involving environmentally-induced rapid directional genetic adaptations make biolog-
ical sense in the context of cosmic Panspermia allowing the efficient spread of living systems and genetic
innovation throughout the Universe. The Hoyle-Wickramasinghe Panspermia paradigm also developed
since the 1970s, unlike strictly geocentric neo-Darwinism provides a cogent biological rationale for the
actual widespread existence of Lamarckian modes of inheritance - it provides its raison d'^
etre. Under a
terrestrially confined neo-Darwinian viewpoint such an association may have been thought spurious in
the past. Our aim is to outline the conceptual links between rapid Lamarckian-based evolutionary
hypermutation processes dependent on reverse transcription-coupled mechanisms among others and
the effective cosmic spread of living systems. For example, a viable, or cryo-preserved, living system
travelling through space in a protective matrix will need of necessity to rapidly adapt and proliferate on
landing in a new cosmic niche. Lamarckian mechanisms thus come to the fore and supersede the slow
(blind and random) genetic processes expected under a traditional neo-Darwinian evolutionary
paradigm.
©2019 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND
license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Contents
Preamble - purpose of this article . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................... 00
1. Summary of terrestrial neo-Darwinism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... ..... 00
*Corresponding author. Centre for Astrobiology, University of Ruhuna, Matara, Sri Lanka.
E-mail address: ejsteele@cyo.edu.au (E.J. Steele).
Contents lists available at ScienceDirect
Progress in Biophysics and Molecular Biology
journal homepage: www.elsevier.com/locate/pbiomolbio
https://doi.org/10.1016/j.pbiomolbio.2019.08.010
0079-6107/©2019 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Progress in Biophysics and Molecular Biology xxx (xxxx) xxx
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
2. The evidence for Lamarck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..................... 00
2.1. The rise of neo-Lamarckian acquired inheritance and the collapse of traditional neo-Darwinian thinking on evolution . . . . . . . . . . . ......... 00
2.2. Environmental stimulation as the directional mutational driver . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
2.3. Role of epigenetic gene targeting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
2.4. Rapid genetic adaptation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
2.5. Penetration of the Weismann Barrier ...................................................... . .. . ... ... ... . .. . .. . .. . ... . .. . .. . .. . .. 00
2.6. Horizontal gene transfer (HGT) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
2.7. Central role of reverse transcription . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
3. Acquired inheritance phenomena . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ............ ......... 00
3.1. Somatic Selection Hypothesis (1979): origin, maintenance, diversification of antibody V genes . . . . . . . . . . . . . . . . . . . . . . . ................ 00
3.2. Inheritance of acquired neonatal tolerance to foreign histocompatibility antigens in mice . . . . . . . . . . . . . . . . . . . . . . . . .................... 00
3.3. The sire effect, telegony and subsequent maternal influence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
3.4. Pavlovian conditioning, coupled maternal influence: brain, behaviour, immunity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ...................... 00
3.5. Transgenerational “epigenetic”experiments in endocrine metabolic systems in rodents . . . . . . . . . . . . . . . . . . . . . . . . . ..................... 00
3.6. The Dutch Famine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
3.7. Uptake of foreign DNA by spermatozoa and inherited effects in progeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
3.8. Adaptive mutations in bacteria and other micro-organisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
3.9. Deaminases, cancer progression and next generation sequencing analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ............................ 00
3.10. Inheritance of characters acquired by plant grafting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
3.11. Complexity of epigenetic and induced transgenerational inheritance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
3.12. 100 years ago? - Experiments of Paul Kammerer, Guyer and Smith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
3.13. Summary and conclusion- Definition of a “gemmule"......................................... . .. . .. . ... ... . .. . .. . .. . ... ... . .. . . 00
4. Pragmatic position: Demarcation Data and the statistical odds of abiogenesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..................... 00
4.1. Eukaryotic and prokaryotic microfossils in carbonaceous meteorites dated at >4.5 billions years . . . . . . . . . . . . . . . . . . . . . ................ 00
4.2. Abiogenesis: life arose from non-living chemistry on earth - what are the statistical odds ? . . . . . . . . . . . . . . . . . . . . . . . . . . ................ 00
5. Lamarck and Panspermia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ............ ......... 00
5.1. Panspermia provides the raison d'etre for Lamarckian Inheritance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
5.2. Growing astronomical evidence from interstellar dust and comets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
5.3. Space survivability of microbiota, and habitable planets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
5.4. Transfer of evolved living systems across the galaxy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
5.5. Some quantitative estimates - cosmic distribution and numbers of living systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . ...................... 00
5.5.1. Space Hardiness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
5.5.2. Effective seeding population sizes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
5.5.3. Protection radiation damage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
5.5.4. Cryopreservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
5.5.5. Exoplanets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
5.6. Evidence from the near-earth environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
5.7. Scientific pragmatism: the near earth neighbourhood? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................... 00
5.8. Cosmic octopus? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .................................................. 00
6. Panspermia and Lamarckian inheritance are no longer mere “Hypotheses”.................................... .. ... .. ... ... . .. . .. ... .. ... . 00
Authors' Note............................................................... .. .................................................... 00
Appendix by Robert Temple: On the Panspermia of the Ancients, Cosmic Spermata and Speculation on Birkeland Currents and Mechanisms of Space
Journeys ....................................................................... ... . . .. . . .. . . ... . ... . ... . ... . ... . ... . . .. . . .. . . ... . ... . .... 00
Transport through space of the microbiota . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................................... 00
Dust clouds ................................................................ ................................................... 00
References ................................................................. .. .. . .................................................. 00
Appendix References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .................................................. 00
Preamble - purpose of this article
All of us have contributed these past 50 years to the assembly of
biological, biophysical and astrophysical data consistent with both
Lamarckian modes of evolution and the conclusion that life itself is
not specifically restricted to Earth but is a cosmic phenomenon
(Panspermia). Our purpose then in writing this speculative review
is to assemble the relevant molecular, cellular, evolutionary and
astrobiological data in a coherent new evolutionary synthesis. We
make the scientific case for the efficient spread and further evo-
lution of pre-existing diverse living systems, unicellular or multi-
cellular, prokaryote, archaea and eukaryote, throughout the
observable universe. While we discuss the problems of the odds
against the emergence of life from non-living chemistry on Earth
our pragmatic position leaves the mechanisms for the ultimate
cosmological origin of life in the Universe an open question. We
make this clear at the outset - this article is not about the origin of
life per se. However our article is concerned with how life has been
continuously seeded to Earth from the Cosmos, and how terrestrial
life has evolved as we recently discussed in this journal (Steele
et al., 2018, 2019). Obviously, all our discussions on Lamarckian
modes of inheritance are derived from observations and experi-
ments about living systems in habitats here on Earth. This provides
valuable insight, in our opinion, of how life can spread throughout
the Cosmos and literally infect and colonize every available niche in
which evolution proceeds as a cosmologically defined and con-
nected process. The whole universe is thus a single connected
biosphere.
1. Summary of terrestrial neo-Darwinism
The widely accepted traditional view of the origin of Life and its
further evolution on Earth, in the period after the Hadean Epoch (~4
billion years ago) can be summarised by the following dot-points:
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx2
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
!Life emerged as the first free-living cell on Earth from non-living
chemistry (Abiogenesis), perhaps via an RNA World, 3e4 billion
years ago in one of Darwin's hypothetical “warm little ponds”or
the canonical “primordial soup.”The current consensus is that
hydrothermal deep sea vents are a plausible location for the
origin of terrestrial life (Martin et al., 2008; Baross, 2018; M!
enez
et al., 2018).
!The first primitive free-living cells then replicated and flour-
ished by Darwinian evolution.
!These early cells repeatedly duplicated their genomes and
genes, slowly mutating their DNA sequences and further rear-
ranging their genomes. This allowed progression from the
archaeal and prokaryotic bacterial worlds, thence to diversify
into a vast range of new and diverse cellular species. At the
inception the multiplication and production of viruses by most
cells further aided cell-cell genetic communication.
!All these evolutionary genetic steps arose by random events
which were then preserved by natural selection.
!The emergence of the first free-living eukaryotic cells occurred
by cell-cell fusions and symbiosis events providing the evolu-
tionary path prior to the emergence of the metazoans, multi-
cellular plant and animal life.
!Many of these evolutionary phases were ponderously slow,
taking millions if not hundreds of millions of years. However the
fossil record, as well as phylogenetic nucleic acid and protein
sequence analyses, show that most novel species and life forms
emerge suddenly in a “punctuated”way either persisting to the
present or going to extinction (termed “punctuated equilib-
rium”by Eldridge and Gould).
!Since the explosive events of the Cambrian adaptive radiation
(~542 million years ago), two further major extinctions and
adaptive radiations have been recorded in the fossil record, at
the Permian/Triassic (P/T) boundary (~252 million years ago)
and the Cretaceous/Paleocene (K/T) boundary (~65.5 million
years ago).
!The time intervals between such apocalyptic events, roughly
about every 200e300 hundred million years, suggests a cosmic
orbiting cycle of our Sun and Solar System around the galactic
centre of the Milky Way; and shorter 30 million year cycles as
our star system oscillates through the galactic plane (Clube et al.,
1996, Wickramasinghe, J.T. et a., 2010).
This dot-point summary accurately describes the widely held
scientific view of Life under the umbrella term “neo-Darwinism”,
on which the analytical discipline of “Population Genetics”is firmly
based. This conventional schema is scientifically valuable because
its “big data”statistical methods have allowed the navigation of the
genomes of thousands of diverse organisms made possible by next
generation sequencing. However, we ourselves and many others
over the years have considered neo-Darwinism itself as being in
need of major conceptual reform. While it unquestionably deserves
respect as the over-arching foundation theory of biology it no
longer reflects the actual state of affairs concerning the totality of
life, its history and how it may have emerged and evolved both on
Earth and throughout the Cosmos (Steele, 1979; Hoyle and
Wickramasinghe, 1981, 1982; Bateson et al., 2017; Noble, 2013,
2017, 2019).
Since the 1970s many key lines of scientific investigation have
produced evidence contradicting this comfortable view of Life on
Earth. We shall discuss some of that key evidence below as it per-
tains to inheritance and genetic mechanisms (Sections 2 and 3).
Recently we ourselves and many colleagues have reviewed most of
this salient contradictory evidence in the context of the data sup-
porting Panspermia (Steele et al., 2018, 2019). The clear conclusion
is that the restricted neo-Darwinian view of terrestrial evolution is
untenable and no longer scientifically credible. It is not denied that
evolutionary developments have occurred in the terrestrial setting.
However at key junctures widely accepted observations do not fit
the actually observed data which allows the plausible conclusion
that "…living organisms such as space-resistant and space-hardy
bacteria, viruses, more complex eukaryotic cells, and on very rare
occasions, even fertilized ova and seeds have been continuously
delivered …to Earth so being one important driver of further
terrestrial evolution which has resulted in considerable genetic
diversity and which has led to the emergence of mankind”(Steele
et al., 2018). Thus life on Earth in all its astonishing variety ap-
pears to have been seeded from the wider Cosmos with the further
terrestrial evolution of these space-derived “varieties”occurring
over hundreds of millions and billions of years on Earth (marked, as
it were, by major cosmic bolide “seeding”events caused by passing
star systems and the passage of our solar system through giant
molecular clouds (e.g. see Hoyle and Wickramasinghe, 1993,
Wickramasinghe, J.T. et al., 2010). We will return to this evidence
and the critical arguments in Sections 4,5 and 6.
2. The evidence for Lamarck
2.1. The rise of neo-Lamarckian acquired inheritance and the
collapse of traditional neo-Darwinian thinking on evolution
A simplified overview of the main evidence gathered since the
1970s is summarised in Table 1. This will be expanded on further in
Section 3. The data collected in this period have firmly established
the validity of the neo-Lamarckian evolutionary paradigm. This 50
year period documents the rise of neo-Lamarckian acquired in-
heritance and the collapse of traditional neo-Darwinian thinking
on evolution. These environmentally-induced cellular and molec-
ular processes can now be considered the primary evolutionary
driver mechanisms for the evolution and ongoing diversification of
life on Earth. We then further argue the case (Section 5) that these
DNA and RNA inheritance mechanisms are likely to be general
throughout the Cosmos (e.g. see Wickramasinghe et al., 2018a) and
will likely operate in the efficient Panspermic dispersal of living
systems throughout the Universe. That is, they allow the immediate
proliferation, rapid adaptation and genetic diversification on
landing of the cosmically-derived organisms surviving impact in
their new cosmic niche.
2.2. Environmental stimulation as the directional mutational driver
Tangible signals from the environment in their broadest sense,
play the key driving role in the origins of “directed”physiological
adaptations and mutations which emerge in the “somatic”body of
the organism. For example, by induced stresses such as pathogen-
inducing innate and adaptive immune responses (deaminase-
mediated mutagenesis at Transcription Bubbles, Fig. 1) but there
are others as discussed in Section 3.
Table 1
Evidence consistent with Lamarckian evolutionary processes.
1 Environmental Stimulation as the Directional Mutational Driver
2 Role of Epigenetic Gene Targeting
3 Rapid Genetic Adaptation
4 Penetration of the Weismann Barrier
5 Horizontal Gene Transfer (HGT)
6 Central Role of Reverse Transcription
The summaries of evidence for Horizontal Gene Transfer phenomena are well
covered at the Wikipedia site https://en.wikipedia.org/wiki/Horizontal_gene_
transfer.
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx 3
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
2.3. Role of epigenetic gene targeting
This environmentally-induced phase “lights up”or “targets”
expressed genes for the adaptive regulation of gene expression in
progeny cells and organisms (Fig. 2). The main epigenetic modifi-
cations are the methylation of targeted cytosines (at CpG sites) via
methyltransferases and their subsequent demethylation of such
sites by AID/APOBEC deaminases and/or TET oxidase enzymes
(reviewed in Guo et al., 2011a,b; Nabel et al., 2012). Demethylations
of this type can thus allow reactivation of gene expression in pre-
viously suppressed genes. During the demethylation process such
genes are vulnerable to cytosine to uracil and 5me cytosine to
thymine deaminase mutations via the AID/APOBEC family of de-
aminases causing C-to-U and 5MeC-to-T primary somatic muta-
tions (at DNA and RNA substrates generated at Transcription
Bubbles (Steele and Lindley, 2017,Fig. 1). This first phase of an
induced adaptive response thus involves “soft”Lamarckian inher-
itance, popularly known as “epigenetic inheritance”as it is
reversible, see Skinner 2015, Skinner et al., 2015 for a recent
comprehensive view (and Fig. 2).
2.4. Rapid genetic adaptation
Evolutionary adaptive change can be very fast and directional -
immediately adaptive to a changing environment within one or
two progeny generations. There is rapid genetic update of genomic
DNA sequences being passed on to progeny organisms. Matic
(2019) has recently reviewed these hypermutation rate strategies
allowing survival of populations of living systems in unpredictable
environments.
2.5. Penetration of the Weismann Barrier
There are numerous instances showing the genetic permeability
of the Soma-Germline Weismann Barrier in higher animals. This
actually requires the resurrection (Liu, 2008; Liu and Li, 2016) of the
ancient idea of Pangenesis employed by Charles Darwin, to para-
phrase Democritus “…. that the seed is formed continuously from
all parts of the body”. The published work now describes the
molecular-vesicle variety of Darwin's “gemmules". There is now a
solid foundation for the concept of Pangenesis as a molecular,
cellular and physiological explanation for the inheritance of
environmentally-induced acquired characters in higher animals
(with a Weismann Barrier) and plants (with no traditional Weis-
mann Barrier), see Fig. 2 and legend, and the review by Noble
(2019).
2.6. Horizontal gene transfer (HGT)
Facilitating these genetic diversification and adaptation pro-
cesses are the ubiquitous phenomena associated with horizontal
gene transfer (HGT) involvinggenetic exchanges between cells (and
their viruses) involving all levels of life, prokaryote, archaea,
eukaryote.
2.7. Central role of reverse transcription
Apart from horizontal gene transfer, the next widespread and
mutagenic phase is the vertical transmissions of Lamarckian acquired
adaptations which involves somatic mutation, somatic selection
Fig. 1. The key features of the reverse transcriptase mechanism of somatic
hypermutation (SHM) at Transcription Bubbles - a type of representation of the
deaminase-based “Universal Mutator”likely to operate in many kingdoms of life
(Lindley, 2018; Krishnan et al., 2018). Some elements of this figure have appeared
before, and this figure is a modified combination of parts from (Steele, 2017), Lindley
and Steele (2013), as well as from mechanism figures in Steele (2009, 2016a) and Steele
and Lindley (2017). This is also an adaptation of the target site reverse transcription
process reported in Luan et al. (1993). Shown is an RNA Polymerase II-generated
Transcription Bubble with C-site and A-site substrate deamination events by AID,
APOBEC and ADAR deaminase enzymes, which generates the strand-biased transition
mutation signatures - A-to-G, G-to-A, G-to-T, and G-to-C. DNA strands shown by black
lines; pre-mRNA as red lines; cDNA strands as thick blue lines due to DNA polymerase
h
acting in its reverse transcriptase mode (Franklin et al., 2004; the RT activity of DNA
Polymerase
h
has been independently confirmed recently by Su et al., 2019). Green
bars are Inosines. Shown also is the action of the RNA exosome (Basu et al., 2011)
allowing access of AID deaminase to cytosines on the transcribed strand (TS). The
ssDNA regions on the displaced non-transcribed strand (NTS) are established targets of
AID action. Note that DNA mutations are first introduced as AID/APOBEC-mediated C-
to-U, followed by excision of uracils by DNA glycosylase (UNG), which creates Abasic
sites in the TS (these can mature into single strand nicks with 30-OH ends via the action
of AP endonuclease, Zanotti et al., 2019). These template Uracil and Abasic sites can be
copied into pre-mRNA by RNA Pol II generating G-to-A and G-to-C modifications as
shown (Kuraoka et al., 2003). Following target site reverse transcription (Luan et al.,
1993), this results in G-to-A and G-to-C mutations in the NTS, in a strand biased
manner. Separately at WA targets in nascent dsRNA substrates, adenosine-to-inosine
(A-to-I) RNA editing events, mediated by ADAR1 deaminase, are copied back into
DNA by reverse transcription via Pol-
h
(Franklin et al., 2004; Steele et al., 2006 ). In
theory, ADARs can also deaminate the RNA and DNA moieties in the RNA: DNA hybrid
(Zheng et al., 2017; Steele and Lindley, 2017). The strand invasion and integration of the
newly synthesized cDNA transcribed strand, as well as random-template mismatch
repair (MacPhee, 1995) are hypothesized additional steps (not shown here). In short,
RNA Pol II introduces modifications in the Ig pre-mRNA as it copies the TS DNA with
AID/APOBEC lesions (Uracils, Abasic sites) and this is coupled to A-to-I editing in
dsRNA stem-loops near the transcription bubble (Steele et al., 2006) as well as in
RNA:DNA hybrids within the bubble (Steele and Lindley, 2017). Next, a RT-priming
substrate is formed when the nicked TS strand with an exposed 30-OH end anneals
with the base modified pre-mRNA copying template allowing cDNA synthesis by Y
Family translesion DNA polymerase-
h
, now acting in its reverse transcriptase mode
(Franklin et al., 2004). These 30-OH annealed priming sites could arise due to excisions
at previous AID/APOBEC-mediated Abasic sites. Alternatively, they could arise due to
an endonuclease excision associated with the MSH2-MSH6 heterodimer engaging a
U:G mispaired lesion (Wilson et al., 2005; Zanotti et al., 2019). Shown is an A-to-T
transversion generated at the RT step at a template Inosine. ADAR, Adenosine Deam-
inase that acts on RNA; AP, an Abasic, or apurinic/apyrimidinic, site; APOBEC family,
generic abbreviation for the C-to-U DNA/RNA deaminase family of which AID is a
member (e.g., APOBEC1; APOBEC3 A, B, C, D, F, G, H); AID, activation induced cytidine
deaminase causing C-to-U lesions at WRCY/RGYW C-site motifs in ssDNA; W, A, or U/T;
WA-site, target motif for ADAR deaminase including DNA polymerase-
h
error prone
incorporation in vitro (Rogozin et al., 2001); Y, pyrimidines T/U or C.; R, purine A or G.
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx4
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
and then reverse transcription at the RNA level into germline ge-
nomes of multicellular animals and plants (Steele, 1979). In many
cases this begins via cytosine and adenosine deaminase action
during gene expression - as represented by the key mutagenic
events at Transcription Bubbles as shown in Fig. 1.Thus RNA mod-
ifications brought about by deaminase action are locked into the
genomic DNA by the process of Target Site Reverse Transcription,
TSRT (Fig. 1 and see specifically Luan et al., 1993). The coupling of
the “soft”inheritance of the “epigenetic”first phase with the sec-
ond “hard”germline or DNA inheritance phase, leads to the stable
transmission of the acquired character(s) to cells and progeny or-
ganisms. Lamarckian inheritance can be envisaged therefore as a
two-step process involving Epigenetic-Genetic coupling (Fig. 2).
In Section 3we select representative examples where each one
is a type of conceptual and/or evidential ‘Demarcation Data’point
in its own right - forcing us to choose between the traditional “slow,
random and blind "neo-Darwinian view of Life to the now rapid,
directional and far more accurate Lamarckian-Panspermic coupled
paradigm of biological evolution.
3. Acquired inheritance phenomena
The development of the main conceptual and experimental
steps are outlined here more or less in chronological order as the
field(s) infolded since 1970 when Temin and Baltimore first re-
ported the discovery of reverse transcriptase in RNA tumour viruses
(Temin and Mizutani, 1970; Baltimore, 1970). The implications of
reverse transcriptase for the inheritance of some acquired charac-
ters was made explicit by Howard Temin at this time if not earlier
(Temin, 1970, 1971). These examples are selective and illustrative of
the diversity and thus generality of Lamarckian acquired inheri-
tance phenomena. More detailed technical information and refer-
ences are confined to figure legends for the interested reader to
further explore in depth.
3.1. Somatic Selection Hypothesis (1979): origin, maintenance,
diversification of antibody V genes
The variable (V) genes of higher vertebrate antibodies, or
Fig. 2. Coupled Epigenetic-Genetic Mechanisms of Lamarckian Inheritance with the Penetration of the Weismann Barrier in Higher Animals. The point of the sequence is to
show that genes targeted first for epigenetic transgenerational regulation can mature via a reverse transcription step targeting the same genes causing hard genetic (DNA) in-
heritance. The key epigenetic-genetic concepts are discussed at length in Lindley (2010, 2011) and specifically in Lindley (2018). The properties of cytosine modifications play a key
role in the plasticity of the Epigenetic-Genetic coupling (see Guo et al., 2011a,b, Nabel et al., 2012). The distinction between “soft”and “hard”Lamarckian inheritance is discussed
explicitly in Steele (2016b). The often confused distinction between the Central Dogma of Molecular Biology and the Weismann Barrier (Steele, 1979; 2016b) is discussed and further
clarified in Noble (2018). In the first phase after stimulation, the environmentally-induced “epigenetic”gene regulatory factors (e.g. methylation-demethylation at CpG sites and
other modifications at cytosines; synthesis of small 21 nt-24nt sRNAs such as miRNAs etc. and guided by other non-coding regulatory RNAs) light up and target specific genes and
gene pathways for regulated gene expression. The epigenetic role of long non-coding RNAs (>200 nt) targeting regulatory portions flanking protein-coding genes is covered in
conceptual detail by Mattick (2003, 2018). The recent review by Kulski (2019) shows the functional importance of locus-wide lncRNAs in conserving long ancestral haplotypes at the
human Major Histocompatibility Complex locus - where they act as genomic anchor points for binding transcription factors, enhancers, and chromatin remodeling enzymes thus
regulating transcription and chromatin folding. LncRNAs specifically target DNA sequences usually via RNA-DNA triple helix interactions involving the weaker yet biologically
significant Hoogsteen hydrogen bonding. Hoogsteen base pairing, considerably weaker than Watson-Crick base pairing is varied, in both parallel and anti-parallel configurations
with the RNA sequence aligned in the major groove of the DNA duplex (Li et al., 2016). These allow multiple points of hydrogen bonding over significant sequence lengths (e.g.
Enhancer or Promoter regions) thus allowing gene-specific recognition. RNA-DNA triple helix interactions thus allow targeted delivery of chromatin modifications resulting in either
active transcription (activation via acetyltransferase-associated complexes) or gene silencing (chromatin compaction via methyltransferase -associated complexes). These data and
the analytical methodology are reviewed in detail in Smith et al. (2013, 2017), Buske et al., (2011, 2012) and in Li et al. (2016). Such lncRNA epigenetic regulators are ubiquitously
found in secreted extracellular vesicles and exosomes, particularly in tumours and tumour cell microenvironments (Xie et al., 2019; Chen et al., 2019). Epigenetically “marked”genes
can become targets for AID/APOBEC-deaminase mediated cytosine to uracil (C-to-U) and cytosine to thymine (C-to-T) mutations (at 5Me CpG sites, Morgan et al., 200 4), which
result in G!U and G!T mispairs. Then as shown in part in Fig. 1 these can progress through further error-prone steps of DNA repair following base excision resulting in Abasic sites
and then single stranded DNA nicks in the transcribed strand with 30OH ends that can prime both DNA and RNA-dependent cDNA synthesis (off homologous newly transcribed RNA
sequence templates). These downstream nicks resulting from ncRNA regulatory targeting therefore “open”the DNA in that genomic region to invasion and targeting of previously
base-modified and mutated mRNA sequence templates which can be reverse transcribed and their specific cDNA fragments integrated at these C-sites (and surrounding sequence)
into the genomic DNA, by target site reverse transcription,TSRT (Luan et al., 1993) as discussed at length elsewhere (e.g. Steele, 2016a.Steele and Lindley, 2017) and shown in Fig. 1.
This is a variant of AID/APOBEC deaminase-mediated dysregulated immunoglobulin somatic hypermutation-like responses initiated at C-sites across the cancer genome (Lindley,
2013; Lindley and Steele, 2013; Lindley et al., 2016). ADAR deaminases causing adenosine to inosine modifications in RNA and DNA are also part of this dysregulated Ig SHM-like
response scheme (A-to-I, read out as A-to-G transitions, Lindley, 2013; Lindley and Steele, 2013; Steele and Lindley, 2017). The enzymatic deaminase targeting specific C-sites and A-
sites in DNA and RNA substrates occurs in protein-coding regions in codon context (Lindley, 2013) most plausibly in the 3D environment of stalled Transcription Bubbles (Lindley,
2013; Steele and Lindley, 2017). There is evidence that deamination of 5-methylcytosine (5 mC) and 5-hydroxymethylcytosine (5hmC) and generation of mutagenic C-to-T mu-
tations directly by the activity of AID/APOBEC complexes is an alternative path to successive oxidation reactions by TET enzymes for the initiation and regulation of DNA deme-
thylation (Guo et al., 2011a,b, Nabel et al., 2012, Pastor et al., 2013, Scourzic et al., 2015). The role of extracellular secreted vesicles and exosomes is discussed in Fig. 1 of Steele and
Lloyd (2015) based on the seminal vesicle/exosome data published in Cossetti et al. (2014) (mice) and later in Sharma et al. (2016) (humans). B lymphocytes themselves when
activated by mitogens secrete large numbers of endogenous retroviruses (Moroni and Schumann, 1975; Moroni et al., 1980). The significance of the very high concentrations (>10
11
per ml) of endogenous retroviruses in seminal fluid, surrounding the placenta and actually bound to the heads of spermatozoa (Keissling et al., 1987) challenges the widely held
belief that only one successful sperm affects the internalized genetic cargo at fertilization. The questioning of this commonly held belief is justified given the huge number of
spermatozoa attached to a given ovum. Finally, apoptotic vesicles have also been invoked as DNA/RNA soma-to-germline transmission vehicles (Steele et al., 2002). Indeed there are
many formal similarities between retroviruses and secreted extracellular vesicles (Hoena et al., 2016). The properties of extracellular extruded vesicles and exosomes has been
extensively reviewed (van der Pol et al., 2012) and extracellular membrane vesicles with exported cargos appear across the three domains of life and form a intercellular
communication system (Gill et al., 2018). All this has been recently highlighted by Noble (2019) in the context of Darwin's Pangenesis.
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx 5
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
immunoglobulins (Ig), exist as large arrays in the germline DNA of
very similar V sequences ("50e10 0 V gene segments). In the
germline they are inactive V segments (coding for about 100 amino
acids) but in a mature somatic B lymphocyte in the lymphoid and
blood circulation they rearrange at the DNA level to join with
shorter D and J elements forming transcriptionally active somatic
genes encoding rearranged heavy (VDJ) and light (VJ) chains of the
HL heterodimers of antibody proteins. A viable antigen combining
site is formed from a heterodimer of one heavy (H)) and one light
(L) chain, essentially by a combinatorial protein association sorting
process in any given B cell. In this “somatic configuration”the V[D]J
genes hypermutate following antigenic stimulation. This is a typical
“Darwinian”process of rapid mutation, proliferation of antigen-
selected B cell survivors with large cellular apoptotic death fac-
tors (in so called “Germinal Centres”in peripheral lymphoid organs,
such as spleen, lymph nodes).
Thus many B cells are destined to die (>90%) in Germinal Cen-
tres. The successful antigen-selected mutants, bearing an antigen-
specific receptor on their surface membrane survive to become
affinity-improved, clonally expanded, memory B lymphocytes
(clonal selection). The daughter cells then enter the vascular cir-
culation and seed other lymphoid organs. All the extant in vivo
molecular and cellular evidence indicates the mechanism of Ig
somatic hypermutation (SHM) is driven by antigenic stimulation
via an AID (APOBEC) and ADAR deaminase-coupled Reverse Tran-
scription process (RNA/RT), as shown in outline in Fig. 1 (Steele
et al., 2006; Steele 2016a, 2017;Steele and Lindley, 2017). The
first iteration of the RNA/RT-Ig SHM model was by Steele and
Pollard (1987), and the demonstration that the key error-prone
DNA polymerase-Eta (
h
) involved in Ig SHM is a very efficient
reverse transcriptase was first demonstrated by Franklin et al.
(2004) and independently confirmed recently by Su et al. (2019).
The Somatic Selection Hypothesis (Steele, 1979) was created to
explain the origin, maintenance and diversification of the germline V
gene arrays via the agency of somatic mutation and clonal selection
(Burnet, 1957, 1959) utilizing Temin's harmless endogenous retro-
viruses acting as somatic gene vectors transducing somatic V mutant
sequencesat the mRNA level and shuttling them into the germline of
immunized animals. Rothenfluh (1995) vastly improved the model
by invoking mobile mutant B lymphocytes interpenetrating repro-
ductive tissue and delivering the endogenous V-transducing vectors
more or less directly to germ cells (later apoptotic B cell-derived
vesicles were also invoked as transport vehicles, Steele et al.,
2002). Rothenfluh (1995) also reviewed the evidence that endoge-
nous retroviruses are secreted in large numbers from B lymphocytes
stimulated by antigens and mitogens of foreign pathogens (Moroni
and Schumann. 1975; Moroni et al., 1980).
The Somatic Selection Hypothesis was the first attempt, post the
Lysenko era, to build a viable Lamarckian genetic model for the
penetration of the Weismann Barrier - that was at the same time
consistent with all the known facts of development, molecular
genetics, virology and Mendelian inheritance. As far as the genetic
structure of higher vertebrate germline V gene arrays are con-
cerned it is still the most economical explanation for the origin,
maintenance and further diversification of all the current published
germline V segment data (Steele and Lindley, 2018) which always
bear the hallmark signatures of intense somatic mutation and se-
lection implying regular soma-to- germline V gene feedback during
life and across generations (Blanden et al., 1998;Steele et al., 1998;
Steele and Lloyd, 2015).
3.2. Inheritance of acquired neonatal tolerance to foreign
histocompatibility antigens in mice
These experiments showed that the deep tolerance of immune
reactivity at the level of cytotoxic T lymphocytes (CTL) measured in
in vitro assay systems which was induced in neonatal male mice,
could, after mating those males as adults to females of the same
inbred stain, be passed on to first and second generation progeny
(appearing in the second generation without exposure to the foreign
histocompatibility (H-2) antigens used to set up specific tolerance in
the original father (Gorczynski and Steele 1980, 1981). In later ex-
periments the specific H-2 tolerance in progeny generations corre-
lated with specific delayed skin graft rejection (Gorczynski et al.,
1983). Experiments in the Brent-Medawar laboratory claimed
these acquired paternal transmission experiments could not be
repeated (Brent et al., 1981). The controversy surrounding these
differences in the two studies initially focused on significant differ-
ences in the way the CTL assay was performed in the Toronto/Canada
and Harrow/UK laboratories. However, the Brent et al. (1981) prog-
eny clearly showed significant numbers of delayed skin graft re-
jectors (Steele, 1981) later confirmed by Gorczynski et al. (1983). In
subsequent follow-up studies the same group again claimed nega-
tive paternal transmission results (Brent et al., 1982) when they set
up a smaller number of breeding males than the original experi-
ments - four neonatally treated males, down from ten in the original
protocols (Gorczynski and Steele, 1980; Brent et al., 1981). This
exposed this work to the further criticism that these investigators
had reduced the odds of observing paternal transmission given that
previous observations had shown that only ~ two out of ten neona-
tally H-2 tolerant males routinely documented high transmission of
H-2 specific hyporesponsiveness (Gorczynski and Steele, 1980).
Indeed, Mullbacher and colleagues, conducting breeding experi-
ments at the same time and in the same laboratory as Brent et al.,
with inactive Bebaru virus antigens, demonstrated a positive non-
antigen specific paternal transmission of induced neonatal hypo-
responsiveness at the CTL level with in vitro cytotoxicity assays
(Mullbacher et al. (1983). Later, positive paternal transmission was
demonstrated in experiments in inbred mice to foreign (rat) eryth-
rocytes, using both repeated high dose neonatal male tolerance
(Steele et al., 1984) or single shot immunity to the erythrocytes in
adult males prior to breeding to normal females (Steele, 1984). So
these acquired inheritance immune system effects induced in male
inbred mice were certainly controversial 30e40 years ago, but they
were real, yet complex with respect to mechanism, involving most
likely, in retrospect, all the epigenetic and genetic dimensions
summarised in Fig. 2.
3.3. The sire effect, telegony and subsequent maternal influence
The acquired inheritance phenomena and history associated with
what is called the “Sire Effect”have been reviewed (Lindley, 2010 pp.
22e29). The phenomenon was reported by Gorczynski et al. (1983)
when they tested the normal inbred female mice who had raised
offspring to male mice of the same inbred strain made neonatally
tolerant to repeated doses of foreign lymphoid cells expressing
specific H-2 histocompatibility antigens as just described
(Gorczynski and Steele, 1980,1981). When these mothers were bred
to normal males of the same inbred strain they produced tolerant or
hyporesponsive progeny to the same H-2 antigens as used in the
original neonatal tolerance regime in the original breeding male.
This was a surprising result. Such mothers also passed on the
effect when fostering normal pups, identifying causal factors in the
colostrum and milk (Gorczynski et al., 1983). This is a striking and
important result with implications resurrecting the old observa-
tions surrounding the non-Mendelian breeding results caused by
male sperm and thus phenomena associated with “Telegony”
(Watson et al., 1983). The “Sire Effect”thus opens a Pandora's Box
with wide implications for pure-line animal breeding and wider
societal implications (Lindley, 2010 pp. 22).
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx6
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
The “Sire Effect”, whatever the detailed transmission mecha-
nism, has real-world practical implications (Lindley, 2010
pp.26e27). Wild rabbits in rural Australia were in plague pro-
portions in the 1940s and 1950s causing great damage to agricul-
ture, particularly the sheep and cattle industries. To control these
wild rabbit populations the Commonwealth Scientific and Indus-
trial Research Organization (CSIRO) released rabbits infected with
lethal Myxomatosis virus. The virus was very effective initially in
controlling wild rabbit numbers. But the speed with which
immunological resistance developed caused further investigations.
At first sight it seemed much faster than simple “Darwinian”re-
covery of a resistant residual population after such a large kill
("90%). Indeed careful follow up controlled breeding experimental
work by Bill Sobey and Dorothy Conolly discovered a significant
factor in the rapid spread of resistance (Sobey and Conolly, 1986):
bucks which had recovered from Myxomatosis virus when mated to
a doe who had not previously been exposed to Myxomatosis virus
produced litters that were resistant to the lethal effects of the virus-
a clear paternal transmission or “Sire Effect”as described by
Gorczynski et al. (1983). Thus when a non-immune buck was mated
with a doe that had previously been mated to an immune buck a
significant number of progeny were born with immunity to
Myxomatosis virus. Sobey and Conolly concluded that an unknown
factor transmitted via the semen of the Myxomatosis virus recov-
ered bucks to the normal females which could be further trans-
ferred in other matings to normal non-exposed males.
The molecular-cellular mechanisms of the Myxomatosis virus
sire effect in rabbits, as well as the H-2 antigen-specific maternal
influence in the mother's milk acquired by the mother from the
original neonatally tolerant male have not been analysed. However
obvious candidates for study can be drawn from an array of
epigenetic and genetic transmission effects discussed above and in
Fig. 2. For example they could be related to the small regulatory
RNA-mediated spermatozoa non-Mendelian inheritance effects
described by Rassoulzadegan and colleagues (Rassoulzadegan et al.,
2006; Kiani et al., 2013; Liebers et al., 2014) and other foreign RNA
and DNA in semen and associated with spermatozoa reported by
Spadafora and colleagues (Lavitrano et al., 1989, Zoraqi and
Spadafora, 1997, Cossetti et al., 2014, Smith and Spadafora, 2005,
Spadafora, 1998, 2008, 2018). These effects are also consistent
with the functional role of sperm-associated RNA reviewed in
Ostermeier et al. (2004). Indeed the oocyte during the fertilization
process has many attached spermatozoa, and it is difficult not to
believe that all the unsuccessful non-fertilizing sperm cells have
not left a functional nucleic acid signature behind in the oocyte.
These could be small regulatory RNAs, specific mRNAs as discussed,
or via genetic information in endogenous somatic retroviruses
(Steele, 1979) attached to sperm heads (Keissling et al., 1987;
Rothenfluh, 1995). Indeed it is hard not to think that the specific
nucleic acid cargoes in the seminal fluid vesicles described by
Cossetti et al. (2014) and Sharma et al. (2016) are also not playing a
functional inheritance role.
3.4. Pavlovian conditioning, coupled maternal influence: brain,
behaviour, immunity
There have been numerous studies published on behavioural
traits associated with Pavlovian conditioned immune phenomena
(Ader and Cohen, 1982, 1993; Moynihan and Ader, 1996). The im-
plications of these experiments and observations are far reaching
for understanding the emergence of specific instincts in higher
animals. In some specific conditioning experiments in mice, in
which cyclophosphamide induced immune suppression was
coupled with saccharin in the drinking water as the conditioning
regime, Gorczynski and colleagues subsequently showed not
merely a saccharin-mediated conditioned recall of immune sup-
pression, as initially described by Ader and Cohen (1982) but the
propagation to progeny through several breeding generations of
that conditioned immunosuppression. Gorczynski and colleagues
localised the transmissible entity by a maternal cross-fostering
design to characteristics of the nursing mother (Gorczynski and
Kennedy, 1987). In other experiments they localised these causal
effects to regulation by factors in the colostrum/foetal-placental
unit modified by conditioning phenomena (Gorczynski, 1992).
This conclusion is the same as the maternal immune factors
transferred in the acquired sire effect described earlier by
Gorczynski et al. (1983).
In a subsequent review Gorczynski et al. (2011) argued “there is
now compelling evidence to suggest that a variety of perceived
environmental “insults”to pregnant females, and even to nursing
females in the post delivery period, in the form of physical,
pathogen-related or emotional stressors, can produce significant
perturbations in the immune responses seen in their offspring.”
The mechanisms involved are ill-understood, but at least in part
may depend upon altered activation of the HPA axis, and of altered
cytokine, neurohormone and neutrotransmitter production within
the CNS. Other data imply an evolutionary balance is struck be-
tween changes in maternal behaviour sacrificing some aspects of
maternal innate immunity at the expense of improved immunity in
offspring. It is now acknowledged that even effects as subtle as an
altered dietary behaviour change in the mother can itself produce
profound changes in the microbiome of both mother and offspring,
and this also can potentially have important implications for sub-
sequent immune development. Since these interactions between
behaviour and immune response potential in mothers and their
offspring are reciprocal in nature, an altered immune activation in
the mother may in turn thus evoke altered behaviour in the
offspring, the “loop”essentially becoming closed.
Indeed at this juncture we can ask: How do instincts arise in
evolution? If one studies the range of these strong and lifesaving
reflex actions in humans and animals, logic implies an ultimate
adaptive Lamarckian cause in ancestors. Strong survival instincts
based on prior learnt fear responses must have arisen in our an-
cestors not by random chance events that were selected, but in a
Lamarckian manner, which were then passed on to their progeny en
masse providing a survival value to the small familial and inter-
breeding groups of mammals in the wild. The recent report by Dias
and Ressler (2014) shows that parental mice subjected to Pavlovian
odour fear conditioning before conception produced progeny
generations with specific behavioural sensitivity to the specific
chemical odour used to condition the parents. Unrelated chemical
odours did not trigger a conditioned fear response. Other breeding
experiments established that these specific acquired transgenera-
tional effects are indeed inherited via parental gametes. Thus, both
direct genomic and indirect epigenetic odorant receptor gene tar-
geting appear to act together to establish what we now recognize as
specific instinctual responses involving odorant receptor genes and
behaviour. This is consistent with a coupling of both “soft”and
“hard”inheritance schemes summarised in Fig. 2.
3.5. Transgenerational “epigenetic”experiments in endocrine
metabolic systems in rodents
The maternally-mediated transgenerational effects just
described in the immune and behavioural systems have tradition-
ally been interpreted under the general “Above the Genes”or “Soft”
acquired inheritance or “Epigenetics”paradigm (Jablonka and
Lamb, 1995; Lindley, 2010; Skinner, 2015). This is indeed the first
phase of environmental stimulation as summarised in Fig. 2.
Definitive induced-transgenerational effects have been described
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx 7
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
in the endocrine and metabolic physiological systems as reviewed
by Campbell and Perkins (1988). The most well known are the
studies showing the paternal and maternal transmission of
chemically-induced acquired diabetes in rodents (Okamoto, 1965;
Goldner and Spergel, 1972; Steele, 1988), passed down many
breeding generations via male and female parents without any
further exposure to the diabetogenic inducing agent (Goldner and
Spergel, 1972). Clearly we are dealing here with a complex inter-
active epigenetic and genetic transmission system. In these studies
the strategies described above (Gorczynski et al., 1983) of testing
the potential of mothers mated to affected males for in utero/foetal
effects by subsequent mating of such mothers to normal males or
cross fostering effects via the colostrum and milk were not
conducted.
3.6. The Dutch Famine
The epigenetic transgenerational phenomena described are not
just of academic interest but impact human health. The famous
after effects of the extreme starvation episodes at the end of World
War II, the “Dutch Famine”, underscore the long term inherited
effects (Painter et al., 2008). Indeed Pembrey and colleagues have
reviewed the induction by environmental conditions of grand
parents and parents, such as nutritional deprivation, exposure to
endocrine disruptors, and traumatic stresses, which can lead to
disease susceptibility and altered immunity in the progeny and
descendants (Pembrey et al., 2014).
Isabell Mansuy and colleagues have shown in mouse models of
maternally-induced stress that unpredictable maternal separation
combined with unpredictable maternal stress (MSUS) can lead to
multiple effects in offspring transmitted via the male line up to
three generations. The progeny phenotypes include depressive-like
syndromes, aberrant social recognition, glucose (insulin) dysregu-
lation and deficits in memory. In recent studies their results
demonstrate both metabolic and behavioural symptoms in progeny
mice into the 4th generation. They conclude that their MSUS
induced transgenerational model produces solid and reproducible
transmission effects initiated in the mother of early life adversity of
male offspring (van Steenwyk et al., 2018).
In all these cases in humans and rodents the phenomena are
interpreted as phase 1 “soft”epigenetic effects (Fig. 2) and thus
potentially reversible via epigenetic reprogramming effects. It is
difficult not to think, given the transmission through to four gen-
erations, that there are no associated “hard”DNA changes, related
to the regulated expression of the relevant targeted gene pathways
as outlined and predicted in phase 2 in Fig. 2. It has not escaped our
notice that the phase 1 to phase 2 or soft-to-hard”acquired in-
heritance outlined in Fig. 2 has similarities to the simulated La-
marckian process described earlier (in 1896) referred to as the
“Baldwin Effect”(Simpson, 1953).
3.7. Uptake of foreign DNA by spermatozoa and inherited effects in
progeny
Corrado Spadafora and colleagues in Rome from the late 1980s
(Lavitrano et al., 1989) to the present have published a series of
important papers showing sperm uptake of foreign nucleic acid
molecules and transmission of the genetic information to progeny
organisms. Thus mouse spermatozoa clearly can take up foreign
DNA/RNA molecules and express the genetic information in their
progeny organisms. In some cases they show that a LINE-1-derived
reverse transcription step can execute the copying of the RNA into
DNA. In #10% of cases the DNA sequences are integrated into the
germline genome. In most cases the sperm-absorbed DNA/RNA
exists as extrachromosomal episomes which replicate along with
the host somatic cells during development displaying mosaic tissue
expression (see reviews in Smith and Spadafora, 2005; Spadafora,
2008). Recent work in mice by Cossetti et al. (2014) suggests a
role for exosomes vesicles released into the bloodstream from hu-
man tumour xenografts transferring somatic RNA to spermatozoa.
This work clearly shows there is no physical barrier in sper-
matozoa to the uptake of DNA or RNA, although developmental
stages in spermatogenesis may be more susceptible to foreign
nucleic acid uptake (Zoraqi and Spadafora, 1997).
In his most recent review of all his group'sdata Spadafora (2018)
has arrived at an important generalisation:
“I propose that RNA-containing nanovesicles, predominantly
small regulatory RNAs, are released from somatic tissues in the
bloodstream, cross the Weismann Barrier, reach the epididymis, and
are eventually taken up by spermatozoa; henceforth the informa-
tion is delivered to oocytes at fertilization. In the model, a LINE-1-
encoded reverse transcriptase activity, present in spermatozoa and
early embryos, plays a key role in amplifying and propagating these
RNAs as extrachromosomal structures. ".
This is among the most precise descriptions of Darwin's “gem-
mules”in animals published, and is a mode of transfer which is also
utilised in genetic information transfer in plant graft-hybridization,
described below (Liu, 2018) and consistent with the current way we
now view Lamarckian inheritance, Fig. 2.
Finally in 2002 Patrick Fogarty reported a striking result of
simple intraperitoneal injection of DNA into adult male or female
mice. Employing a technique based on P-element transposons and
delivering DNA transgenes intravenously in simple vesicles, Fogarty
has shown that 50% of progeny from such male mice inherit the
gene sequence (Fogarty, 2002). The critical integration event re-
quires a transposase enzyme. Thus non-cellular DNA can readily
transverse the testes tissue barriers, that normally quarantine the
production of sperm, be integrated into the germline and be
transmitted to progeny. This was a clear demonstration of the ge-
netic penetration of the Weismann Barrier in mammals in a La-
marckian mode typical of what may take place in the wild in a now
familiar Horizontal Gene Transfer event.
3.8. Adaptive mutations in bacteria and other micro-organisms
Somewhat separate from the above developments involving the
Weismann Barrier in multi-cellular sexually differentiated verte-
brates, the work in bacteria and other rapidly multiplying unicel-
lular organisms (such as yeast) are less definitive conceptually. This
is because simple Darwinian selection of rare population variants
could never be ruled out. However very challenging demonstra-
tions of substrate-induced adaptive evolution phenomena in bac-
teria and yeast have been described (Cairns et al., 1988; Hall, 1988;
Rosenberg, 2001). The data focus thinking on the possibility of
rapid mutator mechanisms in microorganisms during the station-
ary phase (slower replication) and this somehow increases the odds
of ‘selecting’an adaptive mutant. Thus Cairns et al. (1988) pub-
lished data suggesting “directional”mutation phenomena in bac-
teria. Mutants of E. coli requiring lactose for growth (lac-) can be
“directed”under certain conditions to produce lacþ(wild-type)
revertants if cultured in the presence of lactose. They interpreted
this phenomenon as being consistent with a Lamarckian process of
adaptive evolutionary genetic change and they provided a reverse
transcriptase-coupled mechanism for the inheritance of acquired
characteristics, much like that proposed earlier (Steele and Pollard,
1987) for the somatic hypermutation process summarised and now
updated in Fig. 1.
So their work then raised the unexpected and exciting possi-
bility that environmentally induced, non-random mutator pro-
cesses dependent on a reverse transcriptase step also occur in
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx8
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
bacteria. This was rapidly confirmed a few months later by the
timely report of Lampson et al. (1989) demonstrating reverse
transcriptase activity in E. coli. Temin dubbed them all “retrons”in
bacteria (Temin, 1989) and many different reverse transcriptase
activities have now been described throughout prokaryotes and
archaea (Liu et al., 2002; Guo H et al., 2011, 2014; Paul et al., 2015).
Indeed Radman (1999) had earlier predicted such polymerase en-
zymes of evolutionary change (Radman, 1974) as part of the now
familiar adaptive “SOS response”in bacteria to a range of envi-
ronmental stress signals (Tippin et al., 2004). What is very
intriguing about all these developments is that the Y family DNA
Polymerases that figure prominently in the SOS response are all
error-prone polymerases and are indeed related by their DNA
encoded sequence to the human Y family DNA repair polymerases
eta, kappa, iota (Ohmori et al., 2001) all of which have been shown
to be efficient reverse transcriptases (Franklin et al., 2004). How-
ever none of the other bacterial Y family DNA polymerase members
have yet to be examined for their RT activity.
So all these different RT activities in bacteria associated with
adaptive mutator responses such as the diversity-generating ret-
roelements (DGR) (Guo H et al., 2011, Guo H et al., 2014; Paul et al.,
2015) in bacteriophage, bacterial and archaeal genomes provide a
unity with the RNA/RT process developed for the Ig SHM in the
higher vertebrate and mammalian immune system. Mammalian
systems seem to be employing an ancient hypermutation strategy-
a targeted RNA template-directed and reverse transcriptase-
mediated hypermutation process (estimated minimum age given
archaeal systems - 3e4 billion years on Earth). Guo et al. (2014) also
comment on the conserved protein folds of Ig domains in the DGR
reverse transcriptases: “These observations suggest that DGR target
proteins and antigen receptors may have evolved different solu-
tions to accommodate sequence diversity in the context of Ig folds."
3.9. Deaminases, cancer progression and next generation
sequencing analyses
The summary in Fig. 1 showing deaminase-mediated attack on
DNA and RNA substrates in the context of the Transcription Bubble
(ssDNA, RNA:DNA hybrids and nascent dsRNA stem loops) is very
relevant to understanding the somatic mutator processes in pro-
gressing cancer genomes (Lindley, 2013; Lindley et al., 2016; Steele
and Lindley, 2017). What is intriguing is the deaminase substrate
analysis of the clinically relevant single nucleotide polymorphisms
(SNPs) in the human germline - the OMIM data base (Online
Mendelian Inheritance in Man) - and in the wider dbSNP itself
(Lindley and Hall, 2018). The first point to note is that 30e40% of all
the SNPs occur at deaminase sequence motifs typical of AID, APO-
BEC3G, APOBEC3B and ADAR deaminase action in somatic cells
during Innate and Adaptive Immunity to pathogens (see glossary
legend Fig. 1). The next points are that >99% of the far larger
number of millions of SNPs in the NCBI dbSNP are mild or benign
(not associated with overt inherited diseases) and in the protein-
coding regions are in typical C-sites and A-sites specifically tar-
geted in codon-context as observed in somatic cancer genomes
(Lindley, 2013; Lindley et al., 2016). Thus deaminase -mediated
non-random mutation patterns appear written into human germ-
lines over evolutionary time. Given the specificity of the targeted
somatic mutation (TSM) signatures of the deaminases, and their
coincidence with many common established C-site and A-site
deamination motifs this suggests a causal direct role for the AID/
APOBEC and ADAR deaminases mutating human germlines, trig-
gered perhaps by innate immune responses to viral infections. But
this speculation implies that human germline DNA is not quaran-
tined from such pathogen-driven SHM-like processes. Currently
some APOBEC deaminases are known to have low level expression
in normal human ovary but not testes (Refsland et al., 2010), and
some ADAR isoforms are significantly expressed in normal human
testes (Picardi et al., 2015). The deaminases could be acting directly
on transcribed regions of the human germline; alternatively they
act first in the tissues which are somatically selected and thus
become physiologically “benign”(somatic polymorphic variant),
then followed up by a soma-to-germline feedback step to deliver
portions of the mutated somatic sequences into their homologous
genomic sites in germline loci. It seems most unlikely that viral
pathogens would be allowed, under normal circumstances, to
stimulate dysregulated AID/APOBEC and ADAR deaminases to go on
a“mutator spree”in oocytes or during spermatogenesis. Clearly
future human studies should focus on these questions. The point of
discussing all this is to raise the possibility that the current DNA
sequencing and Bioinformatics technologies will soon provide
definitive answers to a more accurate understanding of the true
origins of human genetic variation.
3.10. Inheritance of characters acquired by plant grafting
Plant grafting is an ancient agricultural and horticultural prac-
tice that combines the shoot (scion) of one plant with the root
system (stock) of another. Historically graft-induced variations
were recorded to occur in ancient China. Charles Darwin (1868)
however was the first to use the term “graft hybridization”. He
noted that the formation of breeding hybrids through plant grafting
between distinct species or varieties (without the intervention of
the sexual organs). Many such cases of “graft hybrids”were
described by Darwin where shoots produced from grafted plants
exhibited a combination of characters of both stock and scion. It
was understandable that he would explain their formation by his
theory of Pangenesis involving transmissible and transported
“gemmules”in the phloem. Darwin's concept of graft hybridization
was supported by Ivan Michurin, Lucien Daniel, Luther Burbank
and many practical breeders. But there has been a reluctance to
accept the existence of graft hybrids among some geneticists (Liu,
2018).
Trofim Lysenko was a keen supporter of the Lamarckian inher-
itance of acquired characters, and demonstrated experimentally
the conversion of spring wheat into winter wheat and vice versa. He
also accepted the existence of graft hybrids, and led large-scale
experiments on graft-induced heritable changes. A German
geneticist, Hans Stubbe, failed to confirm Lysenko's results, thus he
regarded graft hybridization as part of Lysenko's fraud (Hagemann,
2002). However many epigenetic phenomena are now recognized
in plants (reviewed in Sano, 2010) which would be equivalent to the
first “Soft”inheritance phase in Fig. 2. Yet other studies in flax
demonstrate “hard”inheritance following environmentally
induced adaptive DNA changes (Cullis, 1984).
Over the past few decades, the existence of graft hybrids has
been widely documented, and the results are clear and striking and
regularly reproduced by many plant breeders and horticulturists.
For example, Yosita Shinoto, the former president of the Genetics
Society of Japan, a serious scientist and a man of the highest
integrity, claimed to have obtained graft hybrids in eggplant and
confirmed Lysenko's results (Shinoto, 1955). A similar example of
such a graft hybrid is shown in eggplant by Zu and Zhao (1957) (see
Fig. 3). There has also been increasing evidence for graft-induced
heritable changes in pepper and other plants (Ohta, 1991; Taller
et al., 1998). The key to success is the use of the so-called
“mentor-grafting”method invented by Michurin.
Graft hybridization is now mainly explained by horizontal gene
transfer and genetic transformation (Ohta, 1991). This is supported
by recent experimental evidence that DNA and entire nuclear ge-
nomes can be transferred between plant cells (Fuentes et al., 2014;
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx 9
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
Gurdon et al., 2016). In addition, the long-distance transport of
mRNA and small RNAs is also considered to be involved in the
formation of graft hybrids. Indeed plant hybrid transmission of
small regulatory RNAs, mRNAs and other reporter sequences are
standard experimental tools in plant molecular genetics and
development (Ham and Lucas, 2017). We should now add a reverse
transcription step to lock in such transported RNA phloem infor-
mation into the hybrid seed genomes. Liu (2006) proposed that
“the stock (or scion) mRNA molecules being transferred into the
scion (or stock) ethen reverse-transcribed into cDNA that can be
integrated into the genome of the scion's (or stock's) germ cells,
embryonic cells, callus cells, as well as the somatic cells of juvenile
plants eand thus may be the main mechanism for graft
hybridization”.
3.11. Complexity of epigenetic and induced transgenerational
inheritance
Despite more than several decades of studying induced epige-
netic effects in animals and plants there are large areas of ignorance
due to the complexity and variety of the phenomena. A recent
balanced and sceptical review of the field by Panzeri et al. (2016) of
non-coding RNA directed epigenetic regulation of gene expression
is necessary corrective reading, despite the considerable molecular
detail on non-coding RNAs short and long now accrued (see legend
Fig. 2). The large number of “above the genes”biochemical modi-
fications to various regulated chromatin states justifies their
conclusion: "Histone modifications comprise methylation, acety-
lation, acylation, phosphorylation, ubiquitination, sumoylation,
proline isomerization, citrullination, and ADP ribosilation, but also
more recently identified (and less represented) crotonylation,
butyrylation, propionylation, succinylation, malonylation, hydrox-
ylation, formylation, O-GlcNAcylation, and likely many others yet to
be discovered”. Furthermore, while gene silencing by methylation
at CpGs and the actions of small regulatory 21 nt-24nt RNAs are the
best studied transgenerational modes (e.g. see Kiani et al., 2013;
Liebers et al., 2014) there are also clear cases of active gene upre-
gulation by small RNAs (Portnoy et al., 2011). All these varied
findings that are ongoing suggest that protein-coding genes and
intervening genomic regions are targeted first by base sequence
homologies (e.g. in the small miRNA stem loop structures, and see
Fig. 2 legend) which then direct the epigenetic methylation and
demethylation mechanisms (e.g. Bayne and Allshire, 2005; Molnar
et al., 2010; He et al., 2011; Matzke and Mosher, 2014). It is our
considered view, following Panzeri et al. (2016), that current in-
vestigations on the plethora of epigenetic-genetic couplings as
summarised in Fig. 2 are just the tip of the iceberg, and that many
genetic surprizes will be revealed in the research of coming years.
3.12. 100 years ago? - Experiments of Paul Kammerer, Guyer and
Smith
One hundred years ago, both before, during and after the first
world war there were serious attempts to demonstrate Lamarckian
inheritance in higher animals by Paul Kammerer (Koestler, 1971;
Vargas, 2009; Vargas et al., 2017) and by Michael Guyer and Eliz-
abeth Smith at the University of Wisconsin (one of their key papers
is republished with a modern interpretation in Steele, 2016c). In our
view the definitive Lamarckian inheritance experiments in rabbits
by Guyer and Smith in 1918e1924 on the transmission via the male
line (up to 9 breeding generations) of maternal autoantibody-
induced eye defects are on a level with the foundation work in
genetics by Gregor Mendel. Yet these experiments were performed
and reported in an earlier age antithetical to Lamarck. This was the
time of the emergence of Mendel's rediscovery, and the rolling
destructive controversies (Koestler, 1971) around the mid-wife toad
and salamander experiments of Paul Kammerer enow given a
modern interpretation in terms of current epigenetic concepts
(Vargas, 2009; Vargas et al., 2017).
This period also heralded the birth of modern neo-Darwinism
which became, with RA Fisher's statistical-based ‘Population Ge-
netics' with free recombination at and between all loci across the
higher plant and animal genome (Hill, 2014), the dominant genetic
paradigm for biology in the 20th century. We do not have space to
deal fully with the RA Fisher paradigm which tacitly guides all
Population Genetics thinking and analysis (Hill, 2014). However we
point out that the prominent and genome-wide existence of long
“ancestral haplotypes”in man and domestic livestock both struc-
turally and functionally, are profoundly non-Darwinian genetic
phenomena, in contradiction of the main assumptions of RA Fisher's
free-recombination paradigm (Dawkins et al., 1999; Williamson
et al., 2011; Dawkins, 2015; Steele, 2014; Steele and Lloyd, 2015).
The existence of numerous and diverse functional long non-coding
RNAs which display functional conservation (Mattick, 2003, 2018;
Smith et al., 2013, 2017; Li et al., 2016) is consistent with the
prior ancestral haplotype concepts (Steele, 2014) discovered by
Roger Dawkins and colleagues at the Major Histocompatibility
Complex (MHC). Indeed Yurek Kulski, who has spent many years
working on the long MHC haplotypes described by Dawkins et al.
Fig. 3. Graft-induced variations in eggplant, in which 9-leaf eggplant was grafted
onto white eggplant. First row: left: fruit of white eggplant; Right: fruit of 9-leaf
eggplant; Middle: fruit in the stock of the immediate generation. Second row:
Variant fruit in F1 generation. Third-Fifth row: Variant fruits in the F2 generation.
Reproduced with permission from Zu, D.-M., Zhao, Y.-S. (1957) A study on the vege-
tative hybridization of some Solanaceous plants. Scientia Sinica, 6, 889e903. From
Fig. 5 in Liu (2018).
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx10
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
(1999), recently reviewed the evidence for involvement of evolu-
tionary conserved lncRNAs in the ancestral haplotype phenomena
associated with the MHC region (Kulski, 2019). Indeed some of us
have also published the possibility of reverse transcriptase-coupled
generation of lncRNAs defining the origin and regulatory integra-
tion of long ancestral haplotypes (Steele et al., 2011).
3.13. Summary and conclusion- Definition of a “gemmule"
The general conclusion from our review of some of the main
extant relevant data seems clear: the traditional Weismann Barrier,
assumed for many years as the bedrock and protective foundation
pillar of modern neo-Darwinism (the past 100 years at least) is very
permeable to somatic DNAs and RNAs with inherited influences on
subsequent generations. Thus a Darwinian “gemmule”or “pan-
gene”can be defined as a somatically-derived vesicle loaded with a
functional cargo - amongst other molecules (proteins, lipids, tran-
scription factors) - of specific regulatory or specific coding nucleic
acid information (small regulatory RNAs, mRNAs, lncRNAs and even
DNAs). It is satisfying that the conceptual wheel has now come full
circle (Noble, 2019) such that we are now able to coolly accept such
an important conclusion on how extant life may be evolving on
Earth (Fig. 2).
4. Pragmatic position: Demarcation Data and the statistical
odds of abiogenesis
There are now solid grounds for believing in the reality of the
inheritance of acquired characters as a widespread biological pro-
cess, albeit with different molecular and cellular details in different
living systems, whether unicellular or multicellular. There are also
now good scientific grounds for believing in the in-fall of living
systems from space continuously “seeding”various life forms on
Earth over the past 4 billion years (Steele et al., 2018, 2019) and
Section 5. However the question of the actual of origins of life
whether on Earth or the wider Universe is clouded in mystery e
with roots stretching into the deepest depths of cosmic antiquity.
Here we outline, as we have done before (Hoyle and
Wickramasinghe, 1981, 1993, 1999a, Steele et al., 2018), our philo-
sophical or, if you will, our pragmatic position.
4.1. Eukaryotic and prokaryotic microfossils in carbonaceous
meteorites dated at >4.5 billions years
In our view the hard evidence already exists distinguishing
terrestrial neo-Darwinism from the evidence for Cosmic Pan-
spermia (Steele et al., 2019). We refer to this key evidence as the
“Demarcation Data”.The accrual of this and other multifactorial
evidence has been comprehensively covered in successive books
since the 1970s as the data and observations unfolded by Fred
Hoyle and N. Chandra Wickramasinghe (1978a, 1979, 1981, 1985,
1991, 1993, 2000 with references to all peer-reviewed papers). A
recent detailed summary of all the key evidence covering the
terrestrial atmosphere, bacteria and other micro-organisms, the
comets as protective incubators and amplifiers of living systems,
the infra-red (IR) analyses of interstellar dust and cometary ejecta
among other central issues can be found in Wickramasinghe
(2015a, 2015b, 2018).
The “Demarcation Data”, or data that has a defining interpre-
tation, distinguishing neo-Darwinism from Cosmic Panspermia
have been recently highlighted by us (Steele et al., 2019).
A key focus is on the internal structure of carbonaceous mete-
orites showing clear microfossils of both eukaryotic and prokary-
otic organisms. These published observations have been secured
from independently curated and examined carbonaceous meteorites,
dated at >4.5 billion years old. The data have been confirmed by
experts in four well curated and characterised carbonaceous me-
teorites: Murchison (Pflug and Heinz, 1997; Hoover, 2005, 2011),
Murchison, Orgueil, Mighei (Rozanov and Hoover, 2013), Polon-
naruwa (Wallis et al., 2013; Wickramasinghe et al., 2013). Terres-
trial contamination has been ruled out e.g. the scanning analytical
EM technology now allows confirmation that the mineralised fossil
has the same chemical composition as the surrounding matrix in
which it is embedded.
As scientists we must critically evaluate these four different and
independent “experiments of nature “on their own terms.
Eukaryotic fossils with silica-based hard shells, are prominent in
these fossils. Certainly in the Polonnaruwa meteorite
(Wickramasinghe et al., 2013) the frustules of clear diatoms are
evident and not considered to be contaminants (Fig. 4). Striking
eukaryotic microfossils like this are also evident in the other
carbonaceous meteorites (Rozanov and Hoover, 2013). Clearly these
are features of mature cell biology in astrophysical phenomenathat
require a coherent explanation. “They strongly imply that complex
cell-based life, now immortalised as fossils in carbonaceous me-
teorites, pre-dates the age of the Earth (and solar system). An
explanation based on Panspermia seems unavoidable to us.”(Steele
et al., 2019).
4.2. Abiogenesis: life arose from non-living chemistry on earth -
what are the statistical odds ?
The question of the actual origins of life should, in our view, be
considered in terms of the information content of life as we know it
and thus in terms of pragmatic statistical probabilities. Thus the
actual origins of Life with its near infinite information content,
enshrouded in the deep recesses of Cosmic antiquity are, for all
practical purposes, scientifically unknowable. Our view on this is
unabashedly pragmatic, which forces a philosophical position on
priorities on where research funds should be deployed in the sci-
entific investigation of both the origins of life, e.g. laboratory
‘abiogenetic’experiments, and in searches where it might exist or
arise across the Universe.
What are the main claims of Abiogenesis? What are their
weaknesses in supporting a localised chemical origin of life? The
summary in the excellent and comprehensive Wikipedia article
Fig. 4. Eukaryotic microfossil in Carbonaceous meterorite. voidal-shaped ribbed
structure embedded in the rock matrix of the Polonnaruwa carbonaceous meteorite,
Wickramasinghe et al. (2013), see also other chemical analyses in Wallis et al. (2013).
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx 11
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
needs to be read in association with our analysis here (https://en.
wikipedia.org/wiki/Abiogenesis)eit clearly shows that apart
from the type of laboratory experiments in the Miller-Urey tradi-
tion of the 1950s (showing that many organic molecules can be
created by electrical discharge processes that may have existed on
the early Earth) the whole discussion in this area is built entirely on
hypothesis, assumption and speculation with no evidence anywhere
documenting the emergence of a living cell from non-living
chemistry. This is not surprizing on reflection ethe scientific en-
terprise investigating Abiogenesis is built (to quote the Wikipedia
article) on “the prevailing scientific hypothesis …that the transi-
tion from non-living to living entities was not a single event, but an
evolutionary process of increasing complexity that involved mo-
lecular self replication, self assembly, autocatalysis, and the emer-
gence of cell membranes.”There is a great enthusiasm (Martin
et al., 2008; Lane, 2015) and much inspired laboratory and theo-
retical research for the first RNA replicator (Szostak et al., 2001;
McFadden, 2016) that would have flourished in the hypothesized
RNA world some 3e4 billion years ago. Such a brief summary is not
to downgrade the importance of the question of the likelihood of
Abiogenesis. We have to be cautious, re. contaminations by all
pervasive living systems here on Earth (cf. observations on tryp-
tophan abiosynthesis at shallow subterranean levels at deep sea
hydrothermal vents as in Menez et al., 2018 ewhich need critical
evaluation given the known all pervasive existence of the deep hot
microbiological biosphere, Gold, 1992, 1999). And we need to be
aware there have also been some spectacular claims (RNA self
replicators) that turned out to be based on experimental artifacts,so
skepticism and caution are warranted (Litovchick and Szostak,
2008, retracted 2017; and see https://retractionwatch.com/2017/
12/05/definitely-embarrassing-nobel-laureate-retracts-non-
reproducible-paper-nature-journal/).
Omission of key information in science is not helpful in any
investigation. Nor should we turn a blind eye to certain overriding
philosophical and epistemological difficulties that have been
recognized. The philosopher of science Karl Popper (1974)
expressed a huge problem for abiogenesis theories thus:
“What makes the origin of life and of the genetic code a dis-
turbing riddle is this: the genetic code is without any biological
function unless it is translated; that is, unless it leads to the syn-
thesis of the proteins whose structure is laid out by the code. But ….
the machinery by which the cell (or at least the non-primitive cell,
which is the only one we know) translates the code consists of at
least fifty macromolecular components which are themselves
coded in the DNA. This constitutes a baffling cycle; a really vicious
circle, it seems, for any attempt to form a model or theory of the
genesis of the genetic code ……Thus we may be faced with the
possibility that the origin of life (like the origin of physics) becomes
an impenetrable barrier to science ……”
Certainly the ‘sin’of omission applies to the Abiogenesis field.
Thus the pragmatic statistical probabilities we discuss below are
rarely mentioned at all in the Abiogenesis research literature nor at
the Abiogenesis Wikipedia site (but see the interesting work by
McFadden, 2016, below).
The paucity of supportive direct scientific evidence for Abio-
genesis is in stark contrast with our own efforts to review here all
the extant concrete and positive evidence demonstrating the re-
ality of Lamarckian modes of inheritance in nature (Sections 2, 3)
and the direct positive biophysical and astrobiological evidence
(Fig. 4, Section 5) consistent with the extraterrestrial origins of life
on Earth (Steele et al., 2018, 2019).
As indicated in our Preamble we are not at all pretending to
provide an explanation for the actual origins of life in the Universe.
We have discussed this again recently (Steele et al., 2018 and
Appendix A in that paper). Many commentators, public and private,
consider we are just shifting the problem e“kicking the can down
the road”- and not solving the actual origin of life itself. However
our pragmatic view is that conventional hypothetical explanations
for Abiogenesis - as an explanation for the emergence of life on
Earth - are mathematically and statistically improbable. Indeed the
odds against successful “Abiogenesis”events popping up all the
time around the Cosmos as implied by regular NASA Press releases
and fuelled by conventional thinking (Walker, 2017) are also not
scientifically supported in any way, and are moreover super-
astronomically improbable. This is based on what we know of the
information content of the simplest minimal cell capable of an in-
dependent self-replicating existence. The odds of bootstrap self-
assembly are formidable (despite the argument that the odds are
sequentially reduced by the emergence of “self replication, self
assembly, autocatalysis, and the emergence of cell membranes.”
Thus the odds against a successful Abiogenesis event are formi-
dable. For the emergence of the first independent free living
bacterial-like cell using a minimal number of essential 256 protein-
coding genes (Mushegian and Koonin, 1996), we can expect one
successfulabiogenic trial in 10
5,120
trials (Hoyle and Wickramasinghe,
1999a) an improbable event anywhere in the currently known uni-
verse, as this number far exceeds by many orders of magnitude the
known atomic and molecular resources of the observable universe
(below). It is thus baffling to continue with the insistence that this
event took place in the infinitesimal locales that could ever have
become available on a primitive Earth. If we invoke an intermediary
RNA world hypothesis as a way of reducing the odds, then the
advocation of a successful “first RNA self-replicator”is also highly
improbable at 4
100
(McFadden, 2016). McFadden is an expert and
concedes the improbabilities as advanced by Hoyle and Wickrama-
singhe based on the “Koonin”number of minimal essential genes.
Thus he considers that 4
100
is an impossible number. He tries to
reduce the oddsby his Quantum computing Life Search model, but he
compounds the problem (in our view) by further assuming highly
improbable intermediate steps. The invocation of improbable inter-
mediate steps is a common feature of all experimental modelling of
Abiogenesis.
The enormity of these super-astronomical numbers we consider
is not often fully appreciated (Hoyle and Wickramasinghe, 1999b).
They can be made somewhat clearer by reference to other large
familiar numbers such as the number electrons, protons, and
neutrons in the known Universe, at 10
80
to 10
90
. The magnitude of
these truly vast improbability factors against abiogenesis is such
that their full impact is not being fully appreciated by the main-
stream scientific community. To repeat ourselves, the figure of
10
5,120
for a minimal number of trials far exceeds by many orders of
magnitude the known molecular as well as probabilistic resources
of the observable universe.
So this is our pragmatic and philosophical position, much like
the pragmatic Copenhagen interpretation in Quantum Mechanics.
Yet we realise that it leads to the following, and to many, unpal-
atable conclusions: Abiogenesis is unlikely anywhere in the known
Universe but would be possible in an “Infinite”Universe or one
approaching infinite size where “Big Bangs”need to be considered as
local space-time expansion-contraction phenomena (which we
termed “rolling Big Bangs”, in Appendix A in Steele et al. (2018).
Indeed a powerful raison d'etre for our continuing insistence to
present an alternative cosmological/panspermic viewpoint is that
every avenue of research in laboratory simulations of localised
abiogenesis have led thus far to an impasse.
Craig Venter's recent success in transplanting a synthetically
manipulated genome in an existing bacterial cell has been hailed by
some as a significant step forward in the quest to create artificial life
de novo (Gibson et al., 2010). This claim, however, is in our view
seriously flawed because what was achieved, using the entire
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx12
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
biochemical machinery of a living cell, was to modify or engineer an
existing genome in a fully functioning living cell. Manifestly this is a
far cry from synthesising life. A digitally modified DNA sequence
alone is a world apart from generating a living bacterium.
To move forward scientifically we suggest we adapt and
embrace a realistic and pragmatic position in the scientific search
for extraterrestrial life. We show below that what is knowable can
be gleaned from the evidence already available from Earth-based
observation and experiment, as well as careful observation and
experiment of relevant astrophysical/biophysical phenomena in
our near-Earth neighbourhood (as discussed in Steele et al., 2018,
2019, and see the data of Allen and DT Wickramasinghe, 1981, DT
Wickramasinghe and Allen, 1983, 1986, NC Wickramasinghe Hoyle,
1998, Wainwright et al., 2015, Grebennikova et al., 2018,
Wickramasinghe et al., 2018b, Shatilovich et al., 2018).
We expand on the implications in Sections 5 and 6 in relation to
the spread of pre-existing living systems throughout the cosmosvia
rapid and directional Lamarckian inheritance.
5. Lamarck and Panspermia
5.1. Panspermia provides the raison d'etre for Lamarckian
Inheritance
Here we strengthen the association between Lamarck and Pan-
spermia that we began to assert a few years ago. To us it is plausible
to consider a strong conceptual link between rapid Lamarckian-
based evolutionary processes dependent on reverse transcription-
coupled mechanisms among others (Wickramasinghe and Steele,
2016) and the effective cosmic spread of living systems via Pan-
spermia. Thus our position is embodied in the answer to the
following key question:
“Why, in contrast to the erroneous fundamental assumptions of
neo-Darwinism, should there be widespread evidence for the ex-
istence on Earth for environmentally-driven (non-random, direc-
tional) Lamarckian modes of inheritance in all the kingdoms of
life?”
Indeed one main purpose in writing this review is to be able to
conclude that HeW Panspermia provides the raison d'etre for the
existence of Lamarckian Inheritance per se; a conclusion quite apart
from any controversial engagement with the limitations of neo-
Darwinism itself. Yet we also agree with the reviewer who made
the following important points:
“..In the manuscript, the rapid adaptation processes that are
inherent in the Lamarckian view of evolution (Section 2, 3) are the
sole conceptual links with Panspermia (Section 5of the manu-
script). The authors provide no additional evidence to strengthen
this correlation. This is a critical aspect that requires a more strin-
gent discussion and conceptual justification.”
Our re-joiner to this type of criticism is this ewe are advancing
a conceptual position based on the critical review of the extant
evidence from the Lamarckian Evolution and Panspermia scientific
fields. The new synthesis thus provides the basis for research
programs in space research. Our conceptual position advances a
hypothesized causal link between “Lamarck”and “Panspermia”
which can advance knowledge in a positive conceptual way, greater
than either one alone. As we discuss below, this allows rational
evaluation of data and speculations of outcomes from current
research programs for the search for extraterrestrial life in the near-
Earth neighbourhood. e.g. orbiting biological laboratories on the
ISS, or other similar platforms, to detect potential space-derived
pathogens, incoming eukaryotic microorganisms, bacteria and vi-
ruses (a research program that can be considered part of the “Hoyle
Shield”,Smith, 2013). .
Further to this, it allows us to understand why adaptive
evolution strategies always involve hypermutation and clonal
diversification (Rosenberg, 2001; Matic, 2019), very much like the
antigen-driven somatic hypermutation process in vertebrate im-
mune responses (Steele, 2016a, 2017). We expect this “survival”
strategy to be implemented by all surviving incoming living sys-
tems from space eso the genetic signature of a surviving popula-
tion of progeny organisms (after an in-fall) will always be one of
hypermutation and adaptive diversity. This is an important insight
that can be applied to the epidemiological and genetic analyses of
all incoming (read “rapidly emerging”) diseases from space (Hoyle
and Wickramasinghe, 1979). And it is an insight that supports our
conclusion that Panspermia provides the raison d'etre for La-
marckian evolution which is independent of any other criticism of
traditional neo-Darwinian theory.
We also try here to makequantitative estimates and predictions,
and discuss how living systems may survive in long term space
journeys. Thus on the basis of available evidence Archaea enmeshed
and protected from lethal radiation in salt crystals may survive
space conditions and be revived after at least 100 million years
(Vreeland et al., 2007). The revival of a spore-forming bacterium
Virgibacillus sp from brine inclusions in halite crystals has been
discovered after 250 million years during which time exposure to
the Earth's natural radioactivity may have delivered radiation doses
exceeding those encountered in the typical transit time between
two exoplanetary systems in space (Satterfield et al., 2005). Perhaps
such long-term survivability of bacteria, their spores and archaea is
not surprising. But what about more complex multicellular ani-
mals? For example, a viable, or cryo-preserved, complex multicel-
lular living system (fertilized egg or plant seed) travelling through
space buried deep within an icy bolide or large meteorite could be
transferred to another habitable planet and will need to rapidly
adapt and proliferate on a landing and thawing in a new cosmic
niche. Within more finely dispersed ejecta arising from an impact
of an asteroid on an inhabited planet, DNA from locally evolved life
could similarly be transferred to other distant habitable planets.
Lamarckian mechanisms of environmentally-driven inherited rapid
adaptation discussed above would come to the fore in such situa-
tions and supersede the infinitesimally slow (blind and random)
genetic processes that are expected under the traditional neo-
Darwinian evolutionary paradigm.
5.2. Growing astronomical evidence from interstellar dust and
comets
With the adventof infrared and ultraviolet astronomy through the
1970's the evidence for organic molecules existing on a vast galactic
scale became incontrovertible (e.g. Hoyle and Wickramasinghe,
2000). There is no easy way by which one could argue that the dis-
coveryof vast quantities of complexorganic moleculesin the universe
could be unconnected with life. As in the case of the Earth the over-
whelming bulk ([99.99 …%) of organic molecules present on a
cosmic scale could most plausibly have a biological connotation.
Although a trend emerged to assert without proof that pre-biotic
chemistry or even pre-biotic chemical evolution was taking place
on an astronomical/cosmological scale, the only correct line of argu-
ment in our view is to accept that biology is a galactic, even cosmo-
logical, process (see references in Wickramasinghe, 2015a; b, 2018).
Biological genetic transfers were clearly taking place over astro-
nomical distance scales. Thus de novo origination of improbable
events for life (Abiogenesis) on the Earth or on other galactic habitats
becomes unnecessary. Panspermia, modulated and augmented by
Lamarckian inheritance processes now becomes an inescapable cos-
mic imperative.
The current biophysical and astrophysical evidence strongly
suggests that the dust grains in the interstellar medium have an
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx 13
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
infrared (IR) absorption spectrum typical of desiccated (freeze-
dried) E. coli bacteria (secured in the laboratory by PhD student
Shirwan Al-Mufti). These data, following age-old standard pro-
cedures in Astronomy, were predicted by Hoyle and Wickrama-
singhe in advance of the interstellar dust observations by DT
Wickramasinghe and DA Allen, is shown in Fig. 5 (from Steele et al.,
2018 Fig. 1 insert). This figure shows the normalised IR extinction
(absorption) flux for two independent data sets. The IR absorption
spectrum in the wavelength range 2.9e4.0 (
m
m) for desiccated
(freeze dried) E. coli bacterial cells (solid line). This is an intricate
and complex IR absorption spectrum of living, albeit dried and
dormant, living cells. The observational data points were secured at
each wavelength indicated for IR electromagnetic radiation emitted
23,000 light years away near the centre of the Milky Way. As this IR
light traverses through clouds of the dust grains it is absorbed in a
similar fashion to the IR absorption by dry E. coli cells in the labo-
ratory experiment on Earth.
The data shown in Fig. 5, was confirmed independently by the
team of Okuda et al. (1990) (and see Fig. 4.3b page 43 in Hoyle and
Wickramasinghe, 1993). The Pearson correlation of this paired
comparison data gives ras 0.9324 for N ¼77 pairs. For Okuda et al.
(1990) the rvalue is 0.9275 for N ¼35 pairs. The Pvalues for both
are <10
&9
. That is, we would expect to see such an exact spectral
match by chance alone in more than one billion similar trials (DT
Wickramasinghe, G Briggs, NC Wickramasinghe, EJ Steele unpub-
lished calculations).
The simplest option is to concede that the dust grains in the
interstellar medium have infrared absorption properties over the
entire continuum from 2.8 to 4.0
m
m that are identical to desiccated
bacterial cells. After 1983 many other spectral features of inter-
stellar dust over other wavebands have also found ready explana-
tions in terms of dust particles of biological origin. We note in
particular the IR absorption spectral matches of larger eukaryotic
cells such as diatoms (algae) for the 8e13
m
m infrared range (e.g.
Hoyle et al., 1982; Hoover et al., 1986; Majeed et al., 1988, and see
Wickramasinghe and Hoyle, 1998).
To the best of our knowledge no artificial modelling of com-
pound organic mixtures will produce such invariant and exact
matches to astronomical data with any reasonable set of assump-
tions. It is difficult therefore to provide an interpretation for these
data that avoids Panspermia.
The idea of biology connected with comets also moved swiftly
from speculation to serious science following the last perihelion
passage of Comet P/Halley in 1986. The first investigation of a
comet in the Space Age (ESA's Giotto mission) thus marked a
turning point in the history of cometary science. A dark organic
comet surface (darker than the darkest coal) was vindicated by the
Giotto photometry. More importantly, in our view, D.T. Wickra-
masinghe and D.A. Allen obtained the first 2e4
m
m spectrum of the
dust from an outburst of the comet on 31st March 1986 (D.T.
Wickramasinghe and Allen, 1986), Fig. 6. This spectrum showed
unequivocal evidence of CeH rotational/vibrational stretching
indicating complex aromatic/aliphatic hydrocarbon structures,
which moreover was consistent with a calculated spectrum for
bacterial dust. Similar data have been published since by others
(e.g. Capaccione et al., 2015).
Another significant correspondence with a putative biology
emerged in the Stardust Mission which captured high speed
cometary dust in blocks of aerogel that were later brought back to
laboratories on the Earth. Amongst the minute fraction of surviving
molecular residues found was the most common of amino acids
Glycine together with a complex mixture of hydrocarbons (Elsila
et al., 2009).
The most recent Rosetta Mission to comet 67P/C-G yielded data
that satisfy consistency checks for biology. Fig. 7 shows a close
consistency between the surface properties of the comet and the
spectrum of a desiccated bacterial sample.
The presence of complex organic molecules including the
building blocks of life in comets is amply confirmed, suggesting
fully-fledged microbial life in comets, or the chemical components
of life (Altwegg et al., 2016) that through coincidence match exactly
the spectra of bacteria. The latter more “conservative”view is that
such molecules could be formed by ion-molecule reactions
including surface chemistry on the surfaces of pre-solar or inter-
stellar grains.
Biological catalytic transformations are of course the most effi-
cient processes by which simple organic and inorganic molecules
can be turned into complex biochemistry of the type seen in
astrophysical phenomena (Steele et al., 2019). Once biology gets
started the conversion of non-biological molecules to biomolecules
(Figs. 5e7) proceeds with an unparalleled efficiency as is indeed
evident from our terrestrial experience. Over 99.9 percent of all the
organic material found on Earth is biologically produced. If biology
is permitted to exist and spread on a galactic/intergalactic scale a
Fig. 5. Comparison of the infrared flux (arbitrary units) from the astronomical
source GC-IRS7 near the galactic centre, with the curve predicted for freeze dried
E. coli cells (Allen D.A. and D T Wickramasinghe, 1981). Also see Wickramasinghe, D.T.
and Allen, D.A. (1983). This is a blow up of the inset in Fig. 1 Steele et al., 2018.
Fig. 6. Comparison of the infrared flux (arbitrary units) from the astronomical
source GC-IRS7 near the galactic centre, with the curve predicted for freeze dried
E. coli cells (Allen D.A. and D T Wickramasinghe 1981). Also see Wickramasinghe, D.T.,
Allen, D.A., 1983. This is a blow up of the inset in Fig. 1 Steele et al., 2018.
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx14
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
similar outcome is to be expected astronomically, consistent with
the data in Figs. 5e7. It is useful to point out that the IR and other
specific spectral signatures of complex and specific biochemical
molecules in the interstellar medium - commonly assumed to be a
rich supply of building blocks for cosmic Abiogenesis events - have
a simple explanation: they are the molecular detritus of dead,
broken and dying cells released into the interstellar medium.
5.3. Space survivability of microbiota, and habitable planets
The requirements for an astronomical source of bacteria-like
cosmic dust and complex organic molecules is (a) the operation
of biological replication in a large class of astronomical bodies, and
(b) the assumption that a non-zero fraction of microbiology so
generated survives the transit between such astronomical habitats,
at any rate between nearest neighbours. Both these pre-conditions
have been established over the past several decades. Survival
properties of bacteria under the most hostile space conditions have
been amply demonstrated both in the laboratory and by means of
direct space experiments. Furthermore, over a hundred billion icy
comets that are known to surround our planetary system (the Oort
cloud of comets) have been convincingly shown to be likely habi-
tats for microbiology emicrobial viability and replication being
accomplished within their radioactively heated interiors
(Wickramasinghe et al., 2012). A fraction of the microbes amplified
in comets are returned into interstellar clouds from which new
stars, comets and planets can form. The feedback loop of cosmic
biology is schematically shown in Fig. 8. In addition to the well-
attested survival attributes of bacteria, particularly of extrem-
ophiles (to which we refer later), it should be stressed here that
only a minuscule survival fraction of interstellar bacteria, ≪10
&20
is
required for every circuit in this loop for panspermic transfers to
the maintained (Hoyle and Wickramasinghe, 2000).
Evidence of bacteria has also been discovered recently in
geological sediments (rocks) that formed 4.1e4.3 by ago during the
Hadean Epoch at a time when the Earth suffered an episode of
heavy bombardment by comets and asteroids (Bell et al., 2015). This
new geological evidence also supports the point of view that
impacting comets brought living entities to the Earth, and by
extension similar impacts on other planetary bodies could establish
life elsewhere in the galaxy.
Perhaps most relevant to the ideas of Panspermia and
Lamarckian inheritance are the recent discoveries of habitable
exoplanets occupying the so-called “Goldilocks zone”. The Orbiting
Kepler telescope launched in 2009 has to date reported the dis-
covery of over 3000 exoplanets in a small sampling volume of the
galaxy. Extrapolating from these discoveries the current estimate of
the total number of habitable planets in the galaxy exceeds 100
billion eapproximately one habitable planet for every sun-like star
(Kopparapu, 2013). The estimated mean separation between such
planets can be estimated to be a few light years. In view of all such
recent discoveries it will be foolish to maintain a pre-Copernican
idea that biology is necessarily confined to our planet, and more
importantly that it originated here against manifestly impossible
odds.
5.4. Transfer of evolved living systems across the galaxy
Whilst amplification of microorganisms within primordial
comets could supply a steady source of primitive life (archaea,
bacteria, unicellular eukaryotes and their viruses) to interstellar
clouds and thence to new planetary systems via comets, the genetic
products of evolved life could also be disseminated on a galaxy-
wide scale. Transportation of entire ecologies of evolved aquatic
life on much rarer occasions could also be possible if life-laden
watery worlds (or large frozen fragments thereof) could occasion-
ally collide with new habitats in the “Goldilocks zones”of stellar
systems. It is tempting to speculate that the Cambrian explosion of
“adaptive radiation”on a grand scale was indeed the product of
such a cosmic seeding of “life-laden watery worlds (or large frozen
planetoid fragments thereof)”as discussed (Steele et al., 2018).
Our present-day solar system with its extended halo of ~100
billion comets (the Oort Cloud) moves around the centre of the
galaxy with a period of 240My. Every 40 million years, on the
average, this comet cloud becomes perturbed due to the close
passage of an interstellar molecular cloud (e.g. the Orion Nebula).
Gravitational interaction then leads to hundreds of comets from the
Oort Cloud being injected into the inner planetary system, some to
collide with the Earth. Such collisions can not only cause
Fig. 7. The surface reflectivity spectra of comet 67P/C-G (left panel, Capaccione et al.,
2015; taken at different times) compared with the transmittance spectrum of desic-
cated E-coli (cf. Figs. 5 and 6) over approximately the same wavelength range for
comparison. Also from Wickramasinghe et al., 2018a,b).
Fig. 8. Amplification cycle of cosmic life. Within our galaxy alone about 100 billion
circuits have been completed, one for every sun-like star.
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx 15
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
extinctions of species (as one impact surely did 65 million years
ago, killing the dinosaurs), but they could also result in the expul-
sion of surface material back into space. A fraction of the Earth-
debris so expelled survives shock-heating and could be laden
with viable microbial ecologies of all types as well as genes and
viruses of evolved life. Such life-bearing material could reach newly
forming planetary systems in the passing molecular cloud within a
few hundred million years of the ejection event. A fledgling plan-
etary system thus comes to be infected with terrestrial microbes -
terrestrial genes that can contribute, via horizontal gene transfer, to
an ongoing process of local biological evolution. If every life-
bearing planet transfers genes (bacteria, viruses, somatic cells and
in rare instances deep frozen seeds and even fertilized eggs) in this
way to more than one other planetary system, life throughout the
galaxy on this picture will inevitably constitute a single connected
biosphere.
5.5. Some quantitative estimates - cosmic distribution and numbers
of living systems
There are thus several key factors to consider in the largely
plausible yet speculative scenarios below on the Panspermic
dispersal of living systems. All are based on known hard data (Hoyle
and Wickramasinghe, 1979;1981;Wickramasinghe, 2015a,b) that
have recently been reviewed (Wickramasinghe, 2018; Steele et al.,
2018, 2019).
5.5.1. Space Hardiness
The “space hardy”features of bacteria are legendary, displaying
un-Earthly properties unlikely to be selected for survival on Earth
but certainly in varied space environments (Hoyle and
Wickramasinghe, 1993). A typical example illustrating such resis-
tance properties to radiation, cold, dehydration, vacuum, acid is the
extremophile Deinococcus radiodurans. Thus bacteria and their
spores, other micro-organisms and eukaryotic cells, and some
exemplar microscopic animals (Tardigrades) are by now well
recognized for these space survival properties. For example the
space survival of algae cultures outside the International Space
Station (Leya et al., 2017), of plant seeds (Tepfer and Leach, 2017) as
well as species of bacteria detected by their DNA sequences in the
cosmic dust on the external surface of the ISS (Grebennikova et al.,
2018). This is quite apart from the >100 million year survival times
of bacteria and archaea in terrestrial salt crystals that we have
already discussed (Vreeland et al., 2007; Satterfield et al., 2005).
5.5.2. Effective seeding population sizes
The population size, N, of a living system within an impacting
bolide that has been ejected from another inhabited planetary or
cometary body must be sufficiently large to permit viable transfer.
The transfer could take the form of viral particles, bacteria, spores
and plant seeds. In some instances even fertilized eggs of insects
and higher animals cannot be excluded. When Nis very large "10
6
it is obvious that a 90% or 99% kill on impact still leaves a significant
number of survivors to rapidly proliferate in a Lamarckian manner
in a congenial niche. This obviously automatically applies to
impacting populations of viruses, bacteria, many microorganisms
and their spores, large populations of plant seeds, and even the
highly contentious suggestion (Steele et al., 2018, 2019) of cry-
opreserved fertilized Cephalopod eggs which could arrive on
impact given large N"10
6
(below).
5.5.3. Protection radiation damage
Encasement in a protective matrix or deep burial is important
both during space travel and on impact. While many organisms
display “space hardy”features, long term survival during space
travel (100 million to billions of years) requires a living system to be
buried or cryopreserved within cometary or planetoid bodies. We
have argued elsewhere that such vehicles as comets or larger
planetoids may act as both protective incubators (active amplifiers)
of living systems and post impact allow significant percentages to
survive and proliferate.
5.5.4. Cryopreservation
In this scenario cryopreservation is a key consideration, partic-
ularly for mature complex multicellular differentiated organisms.
We note the recent observations on the viable recovery of nema-
tode worms from 42,000 year old Late Pleistocene Siberian
permafrost (Shatilovich et al., 2018). Such findings need to be
replicated at other locations and with other species. The data we
have currently available leads to the obvious question: if 42,000
years, why not a billion years of cryopreservation? The discovery
relevant to our argument is that soma-to-germline transfer of ge-
netic information (penetration of the Weisman Barrier) has been
recently demonstrated in C. elegans, a nematode (regulatory double
stranded RNA triggering RNA interference phenomena Devanapally
et al., 2015). This result is of potential importance to the Lamarckian
adaptability of thawed nematodes adapting after arrival in a new
cosmic niche.
All these discoveries have been of crucial importance for the
Hoyle-Wickramasinghe Panspermia paradigm which can include
the transportation of cryopreserved plant seeds and animal em-
bryos within protective matrices (e.g. comets, moons and planets)
or minimally their genes encoded in viruses via undisturbed space
travel extending to hundreds of millions if not billions of years is
not just possible but inevitable. Indeed we would not find it beyond
the realm of possibility that populations of mature microscopic life
forms for example Tardigrades and nematode spp. have a wide
cosmic prevalence and can on occasion be transferred between
suitable cosmic habitats. This would involve a process that merits
being called cosmic Lamarckian evolution. We speculated on such a
scenario for the “sudden”emergence of Cephalopods on Earth 275
mya viz. cryopreserved Octopus eggs (Steele et al., 2018). The same
scenario could apply to a whole range of populations of fertilized
insect eggs of many species as well as plant seeds.
5.5.5. Exoplanets
The number of exoplanetary systems possessing orbiting eco-
systems within habitable zones (comets, moons, planets), with
available water, surface or subterranean, will determine the total
tally of potential cosmic habitats that can be infected. The current
estimates of exo-planets in the habitable zones is >10
10
(Kopparapu, 2013) and possibly much larger if we consider all types
of extreme habits, say, >10
11
. The range of the types of terrestrial
microorganisms existing in the “deep hot biosphere”as first
described by Tommy Gold (1992, 1999) is an indication of the
extraordinary range of extreme habitats that can support life.
Indeed in our Solar System alone there may be an unknown
number of extreme habitats in the form of moons and planetoids on
which large populations of sub-surface living systems could exist
(moons such as Europa, Enceladus, are obvious examples).
We are thus led from many different directions to admit a
convergence to the concept of a genuine “Cosmic Gene Pool”based
on common DNA/RNA/Protein biochemistry (Wickramasinghe
et al., 2018a). Indeed the vast variety of living species on Earth
would be a minuscule subset, albeit a significant sub-set, of an
almost infinite pool.
In terms of magnitudes of incidence of living systems
throughout the Cosmos we might be tempted to rank organisms
(see Table 2) as follows based on a known “Earth equivalent”
multiplied by 10
22
as an educated guess of the number of Earth-like
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx16
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
habitats in the universe (100 billion in every galaxy and 100 billion
galaxies).
Thus the potential cumulative incidence of complex orbiting
ecosystems (on planets, moons, comets) around each observable
star in our galaxy alone begins to approach super-astronomical
magnitudes when these are multiplied by the estimated number
of stars in the observable Universe ~ 10
22
. We should stress at this
point that all the numbers listed above are highly conservative
underestimates according to current thinking in cosmology. Even
within the framework of the currently accepted Big Bang model of
the universe (with early inflation), the observable universe would
only be a minute fraction of what would be a very much larger,
initially causally connected, region that inflated and would thus be
out there, out of contact, but within which Lamarckian transfers
would occur.
5.6. Evidence from the near-earth environment
With some 50e100 tonnes of cometary debris entering the
Earth's atmosphere on a daily basis the collection and testing of this
material for signs of life should in principle at least be straightfor-
ward. Some such projects have been carried out from 2001 on-
wards. The first serious project was carried out with the support of
the Indian Space Research Organisation (ISRO) in partnership with
a group of scientists in the UK including one of us (NCW). Samples
of stratospheric aerosols collected using balloon-borne cry-
osamplers were investigated independently in the UK and India
and revealed evidence of microbial life (Harris et al., 2002). A
particularly interesting component of the collected samples was in
the form of 10
m
m clumps that were identified by SEM and fluo-
rescence tests as being viable but not culturable microorganisms.
Fig. 9 shows putative biological entities discovered in stratospheric
samples by electron microscopy; and the left panel of Fig. 9 shows a
clump of putative cocci and a bacillus. The right panel of Fig. 9
shows evidence of viable microorganisms which did not prove to
be culturable.
Because such large aggregates are virtually impossible to loft to
41 km a prima facie case for their extraterrestrial cometary origin
has been made. A similar experiment to that conducted in 20 01 was
repeated by ISRO in 2009 (Shivaji et al., 2009) and 3 new micro-
organisms were discovered, including one named in honour of Fred
Hoyle as Janibacter hoylei.
The genetic similarities of the new stratospheric bacteria to
existing terrestrial genera have been cited by some as an argument
to discount their possible space origin. However, in our view,
terrestrial bacterial genera all have a space origin, so homologies of
the type found are to be expected and do indeed corroborate a
space origin of all bacteria on Earth (Hoyle and Wickramasinghe,
1979;1981). In order to take the matter further, and hopefully
reach a decisive conclusion, further tests of the collected microbial
samples would be desirable. One such test involves the deployment
of a rather rare laboratory resource ea Nanosims machine. This will
determine the isotopic composition of carbon, oxygen and other
constituent elements within the individual bacterial cells, and if the
composition turns out to be non-terrestrial (Tokoro G, Wickrama-
singhe NC, Temple R and colleagues, experiments underway).
Experiments can also be conducted on the International Space
Station (ISS) to sample the zodiacal cometary dust trails through
which the Earth continuously passes in its orbit around the Sun.
Such initial experiments are reported (Grebennikova et al., 2018;
Wickramasinghe et al., 2018b). Bacteria in the cosmic dust have
been detected by standard PCR techniques on the external surface
of the ISS. These are ground breaking experiments and contami-
nation has been ruled out. Uplifting of micro-organisms to
360e400 km seem quite improbable on physical grounds
(Wickramasinghe and Rycroft, 2018). These data offer the promise
that such ISS microbiological phenomena can be confirmed or
refuted by independent teams of scientists. We can imagine ISS
real-time biological laboratories conducting routine genetic ana-
lyses and tissue cultures using the portable Next Generation
Sequencing (NGS) machines now available. The range of microbial
life, prokaryotic and eukaryotic, in the near Earth cosmic environ-
ment can become, as discussed already, part of the proposed “Hoyle
Shield”, predicting potential pandemics from space (Hoyle and
Wickramasinghe, 1979; Smith, 2013).
Such studies may allow confirmation that Darwinian-
Lamarckian evolution takes place not just within a closed
biosphere on Earth but extends over a large and connected volume
of the cosmos (Wickramasinghe et al., 2018a).
5.7. Scientific pragmatism: the near earth neighbourhood?
In our view pragmatic research on extraterrestrial life is now
required. The research budgets directed to the search for extant and
living extra-terrestrial life needs to be far more focused on the near
Earth neighbourhood. The experiments are relativelycheap and can
be definitive and unequivocal. They offer real-time experimenta-
tion in standard biological laboratories. We have discussed some of
the promising data emerging and there are more recent findings.
Microorganisms have been detected by Milton Wainwright and
colleagues in-falling from space at 41 Km in the Stratosphere
(Wainwright et al., 2015). These data were secured in balloon-lofted
experiments and conducted to avoid terrestrial contamination.
They were set up technically to establish that the microorganisms
and other cellular and viral aggregates were observed following in-
fall not by upwelling. Critics may conclude it is all due to terres-
trial contamination, but the data need to be dispassionately eval-
uated in their own terms. Many of the eukaryotic species detected
can be classed as unassigned Acritarch, and appear viable on impact
with the collection medium. The data more readily fit with Pan-
spermia theory (Wainwright et al., 2015).
Fig. 9. Left: a clump of carbonacesous particles resembling cocci and a bacillus
Right: A clump of viable but non-culturable bacteria fluorescing under the application
of a carbocyanine dye which tests for electric potential across cell walls. Harris et al.,
2002.
Table 2
Cosmic distribution and numbers of living systems.
!Viruses eterrestrial number 10
31
10
53
!Bacteria/Archaea - terrestrial number "10
30
10
52
!Single cell eukaryotes - terrestrial number 10
20
-10
30
10
32
e10
52
!Complex Metazoans - terrestrial number "10
20
10
42
!Higher plants, terrestrial number "10
7
species 10
29
!Higher animals, terrestrial number "10
7
species 10
29
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx 17
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
5.8. Cosmic octopus?
We have tried to discuss throughout this article the “Demarca-
tion Data”which allows distinction between conventional terres-
trial neo-Darwinism (Section 1) and the Cosmic Lamarckian -
Panspermia paradigm. Thus there are a plethora of multifactorial
awkward facts and observations, biological and biophysical, which
fit neatly into the Hoyle-Wickramasinghe Panspermia paradigm
but are often puzzling or inexplicable under a pure neo-Darwinian
terrestrial evolution paradigm - anchored to a super-astronomically
improbable and unproven Abiogenesis event producing the first
cell here on Earth about 4 billion years ago. Most of the relevant
problems and contradictions in this viewpoint are covered in our
recent papers (Steele et al., 2018, 2019). In the same vein a similar
set of awkward facts and observations fall neatly under a La-
marckian world view but not so easily under neo-Darwinism.
Thus we note again the recent observations on the viable re-
covery of nematodes from 42,000 year old Late Pleistocene Siberian
permafrost (Shatilovich et al., 2018). Such findings need to be
replicated at other locations and with other species. Nevertheless
the implications under the Hoyle-Wickramasinghe paradigm sug-
gest that the transportation of cryopreserved complex mature an-
imals within protective matrices (e.g. comets, moons and planets)
or minimally their genes encoded in viruses via undisturbed space
travel extending to hundreds of millions if not billions of years is
the favoured”cross infection”mode across the Cosmos. This would
remain true even if survival probabilities remain minuscule ewith
massive death rates during catastrophic ejection events (e.g. comet
collisions), followed by further attrition upon re-entry onto
receiving host planets. The hundreds of billions of habitable planets
in our galaxy alone would make exchanges of mature biological
entities a virtual certainty eno matter how ridiculous such a
proposition might appear at first sight. The situation is similar to
the sowing of seeds in the wind emost of them are lost, but so very
many are the seeds that some among them are destined to survive.
It is in a similar way that mature animals can albeit exceedingly
rarely land, thaw out in a favourable cosmic habitat for growth, and
thus undergo further cosmic Lamarckian evolution.
We speculated on precisely such a scenario for the emergence of
Cephalopods on Earth 275 mya viz. cryopreserved Octopus eggs
(Steele et al., 2018). This possibility has provoked much discussion
and some ridicule in our circle of private discussions. However
there is no logic whatsoever by which it can be excluded given the
current data. The whole point of our discussion was to show a 250
million year gap between nautiloid precursors and squid/octopus.
That is the whole point of the discussion in Steele et al., 2018. All the
phylogenetic analyses points that out-all experts agree on this gap
in the molluscian evolutionary record. But such huge punctuated
equilibrium-type gaps permeate the fossil/phylogenetic emergence
record, not just the molluscian evolutionary record. That is why
Eldridge-Gould is discussed at length eit is a major unexplained
conundrum (see Fig. 6 in our paper Steele et al., 2018).
6. Panspermia and Lamarckian inheritance are no longer
mere “Hypotheses”
It is reasonable to assert that a scientific theory is a “mature
hypothesis”surviving rigorous critical analyses and hard observa-
tion and experiment. However, it is still in essence Popperian and
thus vulnerable. It can, in principle, be refuted or modified from its
original form, as for instance Einstein's modification/generalisation
of Newtonian mechanics. On the other hand, a hypothesis is usually
the first tentative public utterance of a provisional explanation of a
given set of natural phenomena. It will only mature into a “theory”
if it survives refutation by severe demarcation tests involving
further critical analyses, observation and experiment. By these
criteria the modern field of Cosmic Biology/Panspermia, first clearly
advanced by Hoyle and Wickramasinghe, can be deemed a mature
scientific theory. It provides a coherent explanation for both the
origin of life on Earth and its further non-linear progress of
terrestrial evolution and adaptation as reviewed recently by Steele
et al. (2018).
The HeW thesis has thus survived numerous demarcation tests,
and it has offered many predictions that have been subsequently
fulfilled and furthermore has strong explanatory and predictive
power. For example, a key prediction concerns the distribution and
number of living systems in the known Universe. This distribution
is dictated solely by the “habitability”or otherwise of available
viable Cosmic niches (comets, moons, planets - both orbiting or
wandering) and the DNA/RNA/Protein paradigm for life will hold
across the Cosmos (Wickramasinghe et al., 2018a). This is an
important and definite prediction.
HeW Panspermia theory thus brings together a range of
multifactorial biological facts and phenomena, at first sight unre-
lated, providing a coherent explanation for their existence, their
biological form and their ongoing evolutionary features (Steele
et al., 2018). It therefore provides a general mechanism for the
widely accepted evolutionary pattern of “Punctuated Equilibrium”
described clearly by Eldredge and Gould (1972), then Gould and
Eldredge (1977) but which otherwise remains a semantic descrip-
tion of the known facts. It also predicts and qualifies the ‘genetic’
boundaries of HeW theory. Thus extraterrestrial life is expected, as
just discussed, to possess the same biochemistry, genetic code, the
same DNA and RNA as life on Earth. A radically different life form
discovered would be significant evidence against a universal
Galactic panspermia and the HeW theory would require modifi-
cation (Wickramasinghe et al., 2018a). Yet Galactic-wide pan-
spermia,as we outline here, is now being widely accepted by the
mainstream astronomical community. Thus the Harvard group of
Ginsburg et al. (2018) have recently developed a mathematical
model of Galactic-wide panspermia in which icy comets or rocky
asteroids carrying microbiota could be widely distributed in the
galaxy and exchanged between planetary systems. Their calcula-
tions lend further support to the HeW model of cosmic biology and
our present thesis of Lamarckian transfer in which the galaxy and
the wider universe become a single connected biosphere.
We have also discussed above the vast amount of new evidence
that has accumulated since the 1970s consistent with Lamarckian
Acquired Inheritance phenomena from bacteria through to plants
and animals. Indeed the “Hypothesis of Lamarck”has now survived
some stringent and severe tests earning recognition as a maturing
“Theory”of biological evolution. Our discussion here has focused
where possible on the molecular mechanisms which are now much
clearer in many cases than they were 40 years ago. And our dis-
cussion here has also been on much of the key “Demarcation Evi-
dence”which, as with Cosmic Panspermia, needs to be confronted
by the scientific mainstream. Lamarckian inheritance of acquired
characteristics, based, at their core, on RNA and reverse transcrip-
tase steps, has successfully run the gauntlet of numerous “Pop-
perian”tests during the past 40 years. In our considered opinion the
effective spread of living systems throughout the Cosmos is both by
Lamarckian and Darwinian mechanisms - with the emphasis on
Lamarckian evolutionary processes as these provide rapid and
“directional’adaptations to new Cosmic niches immediately after
the organisms have landed.
Authors' Note
In addition to the well attested mechanisms for particle trans-
port discussed it is worth considering other modes assisting the
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx18
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
panspermic dispersal of living systems throughout the Cosmos.
Appendix by Robert Temple: On the Panspermia of the
Ancients, Cosmic Spermata and Speculation on Birkeland
Currents and Mechanisms of Space Journeys
It is a challenging task to write an appendix to the above paper
which would meet some of the standards we have set for the wide
range of evidence for Lamarckian Panspermia. Although I have
some additional scientific observations to make, I will start with an
update to the situation regarding the Prehistory of Panspermia,
which is a scholarly matter relating to early pre-scientific thinking
by ancient peoples of panspermic notions.
In 2006 I delivered a paper entitled ‘The Prehistory of Pan-
spermia: Astrophysical or Metaphysical?‘, which was published in
2007 (Temple, 2007). In this paper I surveyed proto-panspermia
ideas in a variety of ancient cultures, commencing with the
ancient Egyptians. Amongst the ancient Egyptian texts which I
discussed were the Pyramid Texts, written in what is known as ‘Old
Egyptian’, and which date from the 5th Dynasty (early 25th century
BC to mid 24th century BC) but include much earlier archaic ma-
terial. During 2018, this paper came to the attention of Professor
Joanna Popielska-Grzbowska of the Institute of Mediterranean and
Oriental Cultures, Polish Academy of Sciences, in Warsaw. She is an
expert in Old Egyptian and she informed me that in her studies of
the terminology and concepts embodied in the Pyramid Texts my
comments about them had been confirmed. She also said that she
had found much more evidence in those texts expressing proto-
panspermia ideas. She and I will be producing a joint paper on
this subject during 2019, which will considerably enlarge the dis-
cussion of the prehistory of Panspermia.
I would like also to mention that since my 2007 paper, I have
come to know and appreciate the particular interest of Professor
Otto R€
ossler in the same crucial passage by the ancient Greek
philosopher Anaxagoras (510-428 BC) which I had discussed. He
and I are in frequent contact and it is interesting that we both found
the Anaxagoras passage astonishing, though from two separate,
albeit probably compatible, perspectives. R€
ossler's interest relates
to his seminal work in chaos theory (he is for example the
discoverer of the R€
ossler Attractor), and he did not consider this
passage from the point of view of Panspermia at all. But as his views
on the same passage are so important, I feel that I should mention
them and refer readers to the remarkable volume of scientific
dialogue with him, ‘Chaotic Harmony’(Sanayei and R€
ossler, 2014),
which contains non-mathematical accounts of what he saw in that
passage of Anaxagoras. In the meantime Rossler and I have pub-
lished a joint paper in 2019 entitled ‘Early Einstein Completed’,
concerning the equivalence principle and global-c.
I return now to our present time. There are several aspects of the
Panspermia field which in my opinion have scarcely been touched. I
shall address these in a succession of headings.
Transport through space of the microbiota
The part of this subject which we understand very well is the
dispersal of dust and microbiota into the Earth's atmosphere and its
slow drifting down to the surface, namely the 50e100 tonnes per
day of cosmic dust and debris which reaches the surface of our
planet daily, which is referred to in the main text above. We can
also understand the transport of much of this material by comets
and its dispersal from the comet tails. There is nothing about either
of these processes which is really unusual or indeed unexpected,
once they have been suggested and brought to our attention, as
they first were by Fred Hoyle and Chandra Wickramasinghe in
numerous publications (which has founded the Science of
Panspermia as we now understand it).
The long life of the microbiota, or should we say cosmic sper-
mata (to use the ancient Greek word), is a more than plausible
concept, as witness the main text above. Thus we know from
available evidence that it is possible for cryopreserved cosmic
spermata to survive interplanetary, interstellar, and even inter-
Galactic distances of travel and still be viable entities.
But what is lacking is a full understanding of all the available
means of transport of spermata in those regions where there are no
comets, or where radiation pressure is somehow ineffective. What
then? How do they get around? So far the thinking seems to be of a
slow drifting through space and a tediously ponderous passive
dispersal, mostly by pressure from light rays from stars (radiation
pressure).
My idea is ‘a better design’, and provides for rapid transport
through space of vast quantities of cosmic spermata, so that the
slow part of the transport process would not be across the vast
interstellar and inter-Galactic distances, but would be when the
particles of dust and biota reach a more cluttered region such as our
solar system, where the entire transportation process would
significantly slow down, like a plane coming in for a landing.
Charged dust entering a protoplanetary or planetary environment
with denser plasmas and higher values of magnetic field might be
magnetically braked with charge repulsion rather than collisions
playing a dominant role.
The hypothesis here is that the means of transporting the cos-
mic spermata could be through the moving of sheathed and
internally structured plasmoids within gigantic Birkeland Currents
on a cosmic scale (Peratt, 1992). They would be hurtled through
vast distances at nearly relativistic speeds between star systems
and even between galaxies. These Birkeland Currents would be
super-highways full not only of the charged particles which we
know them to contain but of opportunistic cosmic spermata
hitching a ride. And lest we think this unlikely, we only need to
understand enough about plasma in space to realise that it is al-
ways full of dust particles, not only grains, and frankly plasma is not
particularly bothered about what kind of dust, since essentially ‘any
old dust will do, and if some of it is “alive”, well what the hell, jump
in for the ride’. And the other thing is that all dust particles are
charged, which thus most likely helps lock them in the cosmic
streams of the Currents.
Chandra Wickramasinghe and I are currently exploring the de-
tails of how this process works, and the evidence of the Birkeland
Currents in space. It provides an additional mechanism for mean-
ingful and effective cosmic transport of the cosmic spermata any-
where and everywhere in the Universe.
Dust clouds
We all know that the Universe is full of gigantic dust clouds,
some 100 light years or more across - so full of them in fact that it
might be asked ‘Does God smoke?’and is the Universe really a
smoke-filled room? There is much more to be said about cosmic
dust clouds and their relationship to Panspermia. Chandra Wick-
ramasinghe and I will also be revisiting this question in a future
more detailed investigation (Temple and Wickramasinghe, 2019). A
key factor in what we have to relate is that all cosmic dust clouds are
charged. The charges can be positive or negative, and even if an
entire cloud appears to have zero net charge in total, the internal
structures within the cloud can be so complicated that isolated and
ensheathed regions within the cloud can have opposite charges
which are isolated, and a vast variety of different regions containing
different things. Within the clouds, as within the Birkeland Cur-
rents, pockets of living things can be isolated from barren regions.
The key to understanding this is to understand complex dusty
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx 19
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
plasmas. We should try now to think of cosmic dust clouds not as
just a lot of stuff floating around at random and see them for what
they possibly are, highly structured and immensely complex
genuine entities having a biological provenance. This will all be
further explained in a forthcoming paper.
References
Ader, R., Cohen, N., 1982. Behaviorally conditioned immunosuppression and murine
systemic lupus erythematosus. Science 215, 1534e1536.
Ader, R., Cohen, N., 1993. Psychoneuroimmunology: conditioning stress. Annu. Rev.
Psychol. 44, 53e55.
Allen, D.A., Wickramasinghe, D.T., 1981. Diffuse interstellar absorption bands be-
tween 2.9 and 4.0 mm. Nature 294, 239e240.
Altwegg, K., Balsiger, H., Bar-Nun, A., et al., 2016. Prebiotic chemical eamino acid
and phosphorus in the coma of comet 67P/Churyumov-Gerasimenko. Sci.Adv.
2016 2, e1600285, 27 May.
Baltimore, D., 1970. RNA-dependent DNA polymerase in virions of RNA tumor virus.
Nature 226, 1209e1211.
Baross, J.A., 2018. The rocky road to biomolecules. Nature 564, 42e43. https://doi.
org/10.1038/d41586-018-07262-8.
Basu, U., Meng, F.L., Keim, C., Grinstein, V., Pefanis, E., et al., 2011. The RNA exosome
targets the AID cytidine deaminase to both strands of transcribed duplex DNA
substrates. Cell 144, 353e363. https://doi.org/10.1016/j.cell. 2011.01.001.
In: Bateson, P., Cartwright, N., Dupre, J., Laland, K., Noble, D. (Eds.), 2017. New trends
in evolutionary biology: biological, philosophical and social science perspec-
tives. Interface Focus, 7, 20170 051. https://doi.org/10.1098/rsfs.2017.0051.
Bayne, E.H., Allshire, R.C., 2005. RNA-directed transcriptional gene silencing in
mammals. Trends Genet. 21, 370e373.
Bell, E.A., Boehnke, P., Harrison, T., Mao, W.L., 2015. Potentially biogenic carbon
preserved in a 4.1 billion-year-old zircon. Proc. Natl. Acad. Sci. U.S.A. 112,
14518e14521.
Blanden, R.V., Rothenfluh, H.S., Zylstra, P., Weiller, G.F., Steele, E.J., 1998. The
signature of somatic hypermutation appears to be written into the germline IgV
segment repertoire. Immunol. Rev. 162, 117e132.
Brent, L., Chandler, P., Fiertz, W., Medawar, P.B., Rayfield, L.S., Simpson, E., 1982.
Further studies on supposed Lamarckian inheritance of immunological toler-
ance. Nature 295, 242e244.
Brent, L., Rayfield, L.S., Chandler, P., Fiertz, W., Medawar, P.B., Simpson, E., 1981.
Supposed Lamarckian inheritance of immunological tolerance. Nature 290,
508e512.
Burnet, F.M., 1957. A modification of Jerne's theory of antibody production using the
concept of clonal selection. Aust. J. Sci. 20, 67e69.
Burne, F.M., 1959. Clonal Selection Theory of Acquired Immunity. Cambridge:
Cambridge University Press.
Buske, F.A., Mattick, J.S., Bailey, T.L., 2011. Potential in vivo roles of nucleic acid
triple-helices. RNA Biol. 8, 427e439. PMC3218511.
Buske, F.A., Bauer, D.C., Mattick, J.S., Bailey, T.L., 2012. Triplexator: detecting nucleic
acid triple helices in genomic and transcriptomic data. Genome Res. 22,
1372 e1381. https://doi.org/10.1101/gr.130237.111.
Cairns, J., Overbaugh, J., Miller, S., 1988. The origin of mutants. Nature 335, 142e145.
Campbell, J.H., Perkins, P., 1988. Transgenerational effects of drug and hormonal
treatments in mammals: a review of observations and ideas. In: Boer, G.J.,
Feenstra, M.G.P., Mirmiran, M., Swaab, D.F., van Haaren, F. (Eds.), Progress in
Brain Research, vol. 73, pp. 535e553.
Capaccione, F., Coradini, A., Filacchione, G., Erard, S., Arnoldf, G., et al., 2015.
The organic-rich surface of comet 67P/Churyumov- Gerasimenko as seen by
VIRTIS/Rosetta. Science 347 (6220).
Chen, F., Wang, N., Tan, H.Y., Guo, W., Zhang, C., Feng, Y., 2019. The functional roles
of exosomes-derived long non-coding RNA in human cancer. Cancer Biol. Ther.
20 (5), 583e592. https://doi.org/10.1080/15384047.2018.1564562, 2019.
Clube, S.V.M., Hoyle, F., Napier, W.M., Wickramasinghe, N.C., 1996. Giant comets,
evolution and civilization. Astrophys. Space Sci. 245, 43e60.
Cossetti, C., Lugini, L., Astrologo, L., Saggio, I., Fais, S., Spadafora, C., 2014. Soma-to-
Germline transmission of RNA in mice xenografted with human tumour cells:
possible transport by exosomes. PLoS One 9 (7), e101629. https://doi.org/
10.1371/journal.pone.0101629.
Cullis, C.A., 1984. Environmentally induced DNA changes. In: Pollard, J.W. (Ed.),
Evolutionary Theory: Paths into the Future. John Wiley, London, pp. 217e237.
Darwin, C., 1868. The Variation of Plants and Animals under Domestication. John
Murray, London.
Dawkins, R.L., 2015. Adapting Genetics: Quantal Evolution after Natural Selection e
Surviving the Changes to Come. Nearurban Publishing, Dallas, TX, ISBN 978-0-
9864115-1-9.
Dawkins, R.L., Leelayuwat, C., Gaudieri, S., Tay, G., Hui, J., Cattley, S., et al., 1999.
Genomics of the major histocompatibility complex: haplotypes, duplication,
retroviruses and disease. Immunol. Rev. 167, 275e304.
Devanapally, S., Ravikumar, S., Jose, A.M., 2015. Double-stranded RNA made in C.
elegans neurons can enter the germline and cause transgenerational gene
silencing. Proc. Natl. Acad. Sci. U.S.A. 112, 2133e2138. https://doi.org/10.1073/
pnas. 1423333112.
Dias, B.G., Ressler, K.J., 2014. Parental olfactory experience influences behavior and
neural structure in subsequent generations. Nat. Neurosci. 17, 89e96.
Eldredge, N., Gould, S.J., 1972. Punctuated equilibria : an alternative to phyletic
gradualism. In: Schopf, T.J.M. (Ed.), Models in Paleobiology. Freeman Cooper,
San Francisco, pp. 82e115.
Elsila, J., Glavin, D.P., Dworkin, J.P., 2009. Cometary glycine detected in samples
returned by Stardust. Meteorit. Planet. Sci. 4 4, 1323e1330.
Fogarty, P., 2002. Optimizing the production of animal models for target and lead
validation. Targets 1 (3), 109e116. https://doi.org/10.1016/S1477-3627(02)
02198-0.
Franklin, A., Milburn, P.J., Blanden, R.V., Steele, E.J., 2004. Human DNA polymerase-
h
an A-T mutator in somatic hypermutation of rearranged immunoglobulin
genes, is a reverse transcriptase. Immunol. Cell Biol. 82, 219e225. https://
doi.org/10.1046/j.0818-9641.2004.01221.x.
Fuentes, I., Stegemann, S., Golczyk, H., Karcher, D., Bock, R., 2014. Horizontal
genome transfer as an asexual path to the formation of newspecies. Nature 511,
232e235.
Gibson, D., Glass, J., Lartigue, C., Noskov, V., Chuang, R., et al., 2010. Creation of a
bacterial cell controlled by a chemically synthesized genome. Science 329,
52e56. https://doi.org/10.1126/science.1190719.
Gill, S., Catchpole, R., Forterre, P., 2018. Extracellular membrane vesicles (EVs) in the
three domains of life and beyond, 2018 Nov 21 FEMS Microbiol. Rev.. https://
doi.org/10.1093/femsre/fuy042 ([Epub ahead of print]).
Ginsburg, I., Lingam, M., Loeb, A., 2018. Galactic panspermia. Astrophys. J. Lett. 868,
L12 arXiv:1810.04307.
Gold, T., 1992. The deep hot biosphere. Proc. Natl. Acad. Sci. U.S.A. 89, 6045e6049.
Gold, T., 1999. The Deep Hot Biosphere: the Myth of Fossil Fuels. Copernicus.
Springer Verlag, New York.
Goldner, M.G., Spergel, G., 1972. On the transmission of alloxan diabetes and other
diabetogenic influences. Adv. Metab. Disord. 60, 57e72.
Gorczynski, R.M., 1992. Conditioned stress responses by pregnant and/or lactating
mice reduce immune responses of their offspring after weaning. Brain Behav.
Immmun. 6, 87e95.
Gorczynski, R.M., Kennedy, M., 1987. Behaviour trait associated with conditioned
immunity. Brain Behav. Immun. 1, 72e80.
Gorczynski, R.M., Kennedy, M., MacRae, S., Ciampi, A., 1983. A possible maternal
effect in the abnormal hyporesponsiveness to specific alloantigens in offspring
born to neonatally tolerant fathers. J. Immunol. 131, 1115e1120.
Gorczynski, R.M., Steele, E.J., 1980. Inheritance of acquired immunologic tolerance
to foreign histocompatibility antigens in mice. Proc. Natl. Acad. Sci. U.S.A. 77,
2871e2875.
Gorczynski, R.M., Steele, E.J., 1981. Simultaneous yet independent inheritance of
somatically acquired tolerance to two distinct H-2 antigenic haplotype de-
terminants in mice. Nature 289, 678e681. https://doi.org/10.1038/289678a0.
Gorczynski, L.Y., Gorczynski, C.P., Terzioglu, T., Gorczynski, R.M., 2011. Pre-and post-
natal influences of neurohormonal triggering and behaviour on the immune
response of offspring. Adv. Neuroimmune Biol. 1, 39e51.
Gould, S.J., Eldredge, N., 1977. Punctuated equilibria: the tempo and mode of evo-
lution reconsidered. Paleobiology 3, 115e151.
Grebennikova, T.V., Syroeshkin, A.V., Shubralova, E.V., Eliseeva, O.V., Kostina, L.V.,
et al., 2018. The DNA of bacteria of the world ocean and the Earth in cosmic dust
at the international space station. Sci. World J. 2018, Article ID 7360147, 7 pp
https://doi.org/10.1155/2018/7360147.
Guo, H., Arambula, D., Ghosh, P., Miller, J.F., 2014. Diversity- generating retroele-
ments in phage and bacterial genomes. Microbiol. Spectr. 2 (6) https://doi.org/
10.1128/microbiolspec.MDNA3-0029-2014. MDNA3-0029-2014.
Guo, H., Tse, L.V., Nieh, A.W., Czornyj, E., Williams, S., et al., 2011. Target site
recognition by a diversity-generating retroelement. PLoS Genet. 7 (12),
e1002414 https://doi.org/10.1371/journal.pgen.1002414.
Guo, J.U., Su, Y., Zhong, C., Guo-li Ming, G-l., Song, H., 2011a. Emerging roles of TET
proteins and 5-Hydroxymethylcytosines in active DNA demethylation and
beyond. Cell Cycle 10, 2662e2668.
Guo, J.U., Su, Y., Zhong, C., Ming, G.L., Song, H., 2011b. Hydroxylation of 5- meth-
ylcytosine by TET1 promotes active DNA demethylation in the adult brain. Cell
145 (3), 423e434. https://doi.org/10.1016/j.cell.2011.03.022.
Gurdon, C., Svab, Z., Feng, Y., Kumar, D., Maliga, P., 2016. Cell-to-cell movement of
mitochondria in plants. Proc. Natl. Acad. Sci. U.S.A. 113, 3395e3400.
Hagemann, R., 2002. How did East German genetics avoid Lysenkoism? Trends
Genet. 18, 320e324.
Hall, G.H., 1988. Adaptive evolution that requires multiple spontaneous mutations.
1.Mutations involving an insertion sequence. Genetics120, 887e897.
Ham, B.-K., Lucas, W.J., 2017. Phloem-mobile RNAs as systemic signaling agents.
Annu. Rev. Plant Biol. 68, 173e195, 2017. https://doi.org/10.1146/annurev-
arplant-042916-041139.
Harris, M.J., Wickramasinghe, N.C., Lloyd, D., Narlikar, J.V., Rajaratnam, P., et al.,
2002. Detection of living cells in stratosphere samples. Proc. SPIE 4495, 192.
https://doi.org/10.1117/12.454758.
He, Y.F., Li, B.Z., Li, Z., Liu, P., Wang, Y., et al., 2011. Tet-mediated formation of 5-
carboxylcytosine and its excision by TDG in mammalian DNA. Science 333,
1303e1307. https://doi.org/10.1126/science.1210944.
Hill, W.G., 2014. Applications of population genetics to animal breeding, from
Wright, Fisher and Lush to genomic prediction. Genetics 196, 1e16.
Hoena, N-t.E., Cremera, T., Gallo, R.C., Margolisc, L.B., 2016. Extracellular vesicles and
viruses: are they close relatives? Proc. Natl. Acad. Sci. 113, 9155e9161. www.
pnas.org/cgi/doi/10.1073/pnas.1605146113.
Hoover, R.B., 2005. Microfossils, biominerals, and chemical biomarkers in
E.J. Steele et al. / Progress in Biophysics and Molecular Biology xxx (xxxx) xxx20
Please cite this article as: Steele, E.J et al., Lamarck and Panspermia - On the Efficient Spread of Living Systems Throughout the Cosmos, Progress
in Biophysics and Molecular Biology, https://doi.org/10.1016/j.pbiomolbio.2019.08.010
meteorites. In: Hoover, R.B., Rozanov, A.Y., Paepe (Eds.), Perspectives in Astro-
biology. RR IOS Press, Amsterdam, pp. 43e65.
Hoover, R.B., 2011. Fossils of cyanobacteria in CI1 carbonaceous meteorites: impli-
cations to life on comets, Europa and Enceladus. J. Cosmol. 16, 7070e7111.
Hoover, R.B., Hoyle, F., Wickramasinghe, N.C., Hoover, M., Al-Mufti, S., 1986. Diatoms
on Earth, comets, Europa and in interstellar space. Earth Moon Planets 35,
19e45.
Hoyle, F., Wickramasinghe, N.C., 1978. Life Cloud. J.M. Dent Ltd, London.
Hoyle, F., Wickramasinghe, N.C., 1979. Diseases from Space. J.M. Dent Ltd, London.
Hoyle, F., Wickramasinghe, N.C., 1981. Evolution from Space. J.M. Dent Ltd, London.
Hoyle, F., Wickramasinghe, C., 1982. Why Neo-Darwinism Does Not Work. Univer-
sity College Cardiff Press, ISBN 0 906449 50 2.
Hoyle, F., Wickramasinghe, N.C., 1985. Living Comets. Univ. College, Cardiff Press,
Cardiff.
Hoyle, F., Wickramasinghe, N.C., 1991. The Theory of Cosmic Grains. Kluwer,
Dordrecht.
Hoyle, F., Wickramasinghe, N.C., 1993. Our Place in the Cosmos : the Unfinished
Revolution. J.M. Dent Ltd, London.
Hoyle, F., Wickramasinghe, N.C., Al-Mufti, S., et al., 1982. Infrared spectroscopy over
the 2.9-3.9
m
m waveband in biochemistry and astronomy. Astrophys. Space Sci.
83, 405e409.
Hoyle, F., Wickramasinghe, N.C., 1999a. Panspermia 20 00. Astrophys. Space Sci. 268,
1e17.
Hoyle, F., Wickramasinghe, N.C., 1999b. The Universe and Life: deductions from the
weak anthropic principle. Astrophys. Space Sci. 268, 89e102.
Hoyle, F., Wickramasinghe, N.C., 2000. Astronomical Origins of Life : Steps towards
Panspermia. Reprints from Astrophys. Space Sci 268. Kluwer Academic Pub-
lishers, Dordrecht, The Netherlands, pp. 1e3, 1999.
Jablonka, E., Lamb, M.J., 1995. Epigenetic Inheritance and Evolution:The Lamarckian
Dimension. Oxford University Press, Oxford.
Kopparapu, R.K., 2013. A revised estimate of the occurrence rate of terrestrial
planets in habitable zones around Kepler M-Dwarf. Astrophys. J. 767, L8.
Keissling, A.A., Crowell, R.C., Connell, R.S., 1987. Sperm- associated retroviruses in
the mouse epididymis. Proc. Natl. Acad. Sci. U.S.A. 84, 8667-8571.
Kiani, J., Grandjean, V., Liebers, R., Tuorto, F., Ghanbarian, H., et al., 2013.
RNAemediated epigenetic heredity requires the cytosine methyltransferase
Dnmt2. PLoS Genet. 9 (5) https://doi.org/10.1371/journal.pgen.1003498
e1003498.
Koestler, A., 1971. The Case of the Midwife Toad. Pan Books, London.
Krishnan, A., Iyer, L.M., Holland, S.J., Boehm, T., Aravind, L., 2018. Diversification of
AID/APOBEC-like deaminases in metazoa: multiplicity of clades and wide-
spread roles in immunity. Proc. Natl. Acad. Sci. U.S.A. 115, E3201eE3210.
Kulski, J.K., 2019. Long noncoding RNA HCP5, a hybrid HLA Class I endogenous
retroviral gene: structure, expression, and disease associations. Cells 8 (5), 480.
https://doi.org/10.3390/cells8050480.
Kuraoka, I., Endou, M., Yamaguchi, Y., Wada, Y., Handa, H., Tanaka, K., 2003. Effects
of endogenous DNA base lesions on transcription elongation by mammalian
RNA polymerase II. J. Biol. Chem. 278, 7294e7299. https://doi.org/10.1074/jbc.
M208102200.
Lampson, B.C., Sun, J., Hsu, M.-Y., Yallejo-Ramirez, J., Inouye, S., Inouye, M., 1989.
Reverse transcriptase in a clinical strain of Escherichia coli: production of
branched RNA-linked msDNA. Science 243, 1033e1038.
Lavitrano, M., Camaioni, A., Fazio, V.M., Dolci, S., Farace, M.G., Spadafora, C., 1989.
Sperm cells as vectors for introducing foreign DNA into eggs: genetic trans-
formation of mice. Cell 57, 717e723.
Lane, N., 2015. The Vital Question: Energy, Evolution, and the Origins of Complex
Life. W.W. Norton &Company, London.
Leya, T., et al., 2017. Algae Survive outside Space Station. AlgaeIndus-
tryMagazine.Com, 7th February, 2017. http://www.algaeindustrymagazine._
com/algae-survive-outside-space-station/.
Li, Y., Syed, J., Sugiyama, H., 2016. RNA-DNA triplex formation by long noncoding
RNAs. Cell. Chem. Biol. 23, 1325e1333. https://doi.org/10.1016/
j.chembiol.2016.09.011.
Liebers, R., Rassoulzadegan, M., Lyko, F., 2014. Epigenetic regulation by heritable
RNA. PLoS Genet. 10, e1004296. https://doi.org/10.1371/journal.pgen.1004296.
Lindley, R., 2010. The Soma: How Our Genes Really Work and How that Changes
Everything! CYO Foundation. POD book. CreateSpace, Amazon.com, ISBN
1451525648.
Lindley, R.A., 2011. How evolution occurs: was Lamarck also right? EdgeScience #8,
6e9. https://www.scientificexploration.org/edgescience/8.
Lindley, R.A., 2013. The importance of codon context for understanding the Ig-like
somatic hypermutation strand-biased patterns in TP53 mutations in breast
cancer. Cancer Genet. 206, 222e226. https://doi.org/10.1016/
j.cancergen.2013.05.016.
Lindley, R.A., 2018. A new treaty between disease and evolution - are deaminases the
“universal mutators”responsible for our own evolution? EdgeScience #36,
16e20. http://www.scientificexploration.org/edgescience/edgescience-issue-36.
Lindley, R.A., Hall, N.A., 2018. APOBEC and ADAR deaminases may cause many single
nucleotide polymorphisms curated in the OMIM database. Mutat. Res. Fundam.
Mol. Mech. Mutagen. 810, 33e38. https://doi.org/10.1016/j.mrfmmm.2018.03.
008.
Lindley, R.A., Steele, E.J., 2013. Critical analysis of strand-biased somatic mutation
signatures in TP53 versus Ig genes, in genome-wide data and the etiology of
cancer ISRN Genomics, 2013 Article ID 921418, 18 pages. https://www.hindawi.
com/journals/isrn/2013/921418/.
Lindley, R.A., Humbert, P., Larmer, C., Akmeemana, E.H., Pendlebury, C.R.R., 2016.
Association between targeted somatic mutation (TSM) signatures and HGS-
OvCa progression. Cancer Med. 5, 2629e2640. https://doi.org/10.1002/
cam4.825.
Litovchick, A., Szostak, J.W., 2008. Selection of cyclic peptide aptamers to HCV IRES
RNA using mRNA display. Proc. Natl. Acad. Sci. U.S.A. 105, 15293e15298. https://
doi.org/10.1073/pnas.0805837105.
Liu, Y., 2006. Historical and modern genetics of plant graft hybridization. Adv.
Genet. 56, 101e129. https://doi.org/10.1016/S0065-2660(06)56003-1.
Liu, Y., 2008. A new perspective on Darwin's Pangenesis. Biol. Res. 83, 141e149.
https://doi.org/10.1111/j.1469-185X.2008.00036.x.
Liu, Y., 2018. Darwin's Pangenesis and graft hybridization. Adv. Genet. 102, 27e66.
https://doi.org/10.1016/bs.adgen.2018.05.007.
Liu, M., Deora, R., Doulatov, S.R., Gingery, M., Eiserling, F.A., et al., 2002. Reverse
transcriptase-mediated tropism switching in Bordetella bacteriophage. Science
295, 2091e2094.
Liu, Y., Li, X., 2016. Darwin's Pangenesis as a molecular theory of inherited diseases.
Gene 582, 19e22. https://doi.org/10.1016/j.gene.2016.01.051.
Luan, D.D., Korman, M.H., Jakubczak, J.L., Eichbush, T.H., 1993. Reverse transcription
of R2B mRNA is primed by a nick at the chromosomal target site; A mechanism
for non-LTR retrotransposition. Cell 72, 595e605. PMID: 7679954.
MacPhee, D.G., 1995. Mismatch repair, somatic mutations and the origins of cancer.
Cancer Res. 55, 5489e5492.
Martin, W., Baross, J., Kelley, D., Russell, M.J., 2008. Hydrothermal vents and the
origin of life. Nat. Rev. Microbiol. 6, 805e814. https://doi.org/10.1038/
nrmicro1991.
Matic, I., 2019. Mutation rate heterogeneity increases odds of survival in unpre-
dictable environments. Mol. Cell 75, 421e
425. https://doi.org/10.1016/j.molcel.
2019.06.029.
Mattick, J.S., 2003. Challenging the dogma: the hidden layer of non-protein-coding
RNAs in complex organisms. Bioessays 25, 930e939.
Mattick, J.S., 2018. The State of long non-coding RNA biology. Noncoding RNA 4 (3),
E17. https://doi.org/10.3390/ncrna4030017