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Regulating Information in Molecules: The Convention on Biological Diversity and Digital Sequence Information

Charles Lawson

Griffith University, Australia Abstract

Keywords: Convention on Biological Diversity; genetic resources; access and benefit-sharing; DNA sequences; digital sequence information; information.

Introduction

The United Nations Convention on Biological Diversity (CBD) and its subsequent Nagoya Protocol on Access to Genetic Resources and the Fair and Equitable Sharing of Benefits Arising from their Utilization to the Convention on Biological Diversity (Nagoya Protocol) proposed a framework to conserve biological diversity, sustainably use the components of biodiversity and fairly and equitably share the benefits from utilising genetic resources.1 The basic scheme for fairly and equitably sharing the benefits from utilising genetic resources obliges Contracting Parties to the CBD and Parties to the Nagoya Protocol to consider implementing legislative, administrative and policy measures facilitating access to ‘genetic resources’

within their sovereign control with prior informed consent and mutually agreed terms (known as access and benefit-sharing (ABS)).2 In this context, ‘genetic resources’ are defined as ‘genetic material of actual or potential value’ and ‘genetic material’

as ‘any material of plant, animal, microbial or other origin containing functional units of heredity’.3 In practice, however, the term has a very flexible meaning, and Contracting Parties implementing the CBD may apply the term broadly to include most biological materials and derivatives.4 The Nagoya Protocol extends these obligations to include ‘derivatives’5 and to

1 CBD, Art. 1; Nagoya Protocol, Art. 1.

2 CBD, Art. 15; Nagoya Protocol, Arts. 5 and 6.

3 CBD, Art. 2. See also UNEP/CBD/WG-ABS/7/2, [18] and Annex ([3]).

4 See UNEP/CBD/WG-ABS/9/INF/1. See also UNEP/CBD/COP/3/20, [35]–[37].

5 A derivative is ‘a naturally occurring biochemical compound resulting from the genetic expression or metabolism of biological or genetic resources’: Nagoya Protocol, Art. 2. The interpretation is complicated because the term ‘derivative’ is included in the definition of

‘biotechnology’, and that term is then included in the definition of ‘utilization of genetic resources’ that is engaged in the fair and equitable benefit-sharing of the ABS obligations: Arts. 2, 5.1 and 5.2.

The United Nations Convention on Biological Diversity and its subsequent Nagoya Protocol on Access to Genetic Resources and the Fair and Equitable Sharing of Benefits Arising from their Utilization to the Convention on Biological Diversity provide a framework to conserve biological diversity, sustainably use biodiversity components and fairly and equitably share their benefits. There is unresolved contention about treating information as a derivative of biological materials and a distinct commodity with a value that can be translated into definable benefits. This article addresses whether there is information in DNA sequences, finding that there is causal information but no intentional or semantic information, although the causal contribution remains difficult to determine. This article concludes that caution should be exercised in limiting access to information in DNA through regulation because of the perverse outcomes controlling potential uses and reducing incentives for others to use information in new and innovative ways.

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‘Traditional Knowledge associated with genetic resources’.6 The CBD has attracted 196 Contracting Parties and the Nagoya Protocol 133 Parties. Most ABS schemes rely on a contractual arrangement between a resource holder and the party seeking access to the resource that incorporates the CBD and Nagoya Protocol obligations, including the sharing of monetary and non- monetary benefits.7

Despite almost three decades of operation, concerns remain about the likely potential for the CBD and Nagoya Protocol to deliver significant benefits.8 As a result of these concerns, there has been a resurgent interest in extracting benefits from utilising information associated with genetic resources (e.g., downloading a DNA sequence from a publicly available database) as another source of potentially significant ABS benefits. However, failing to include or extend ABS to genetic information could undermine the existing ABS scheme because information can be utilised without the ABS arrangements and specifically benefit-sharing that apply to physical genetic resources.9 At the CBD and Nagoya Protocol forums, these concerns were captured by the term ‘digital sequence information’ (DSI). This term is a ‘place holder, without prejudice to future consideration of alternative terms’.10 The core of the contention is the ways and merits of treating DSI as a derivative of the materials within the ABS transaction itself, which becomes a distinct commodity with a value that the ABS scheme attempts to translate into definable benefits.11

The purpose of this article is to address the concern that informational language in the context of genetic resources (e.g.,

‘transcription’, ‘translation’, ‘coding’, ‘editing’, ‘proofreading’, ‘copying’, ‘gene expression’, ‘signals’, ‘program’ and ‘book of life’)12 relating to morphological development and evolution potentially falling within the scope of the CBD and the Nagoya Protocol’s ABS arrangements is essentially flawed. While these are not new concerns in the discussion on information metaphors in the biological sciences, they are poignant because a bottom-up account of genetics based on the information flowing from DNA sequences infers a significance for a sequence that it might not have. In contrast, a top-down account traces an observable phenomenon to the products that result from the DNA sequence and other relevant causes. Put simply, as this article will demonstrate, the informational language used to describe molecular biology13 ‘leads to a misleading picture of possible explanations in molecular biology’14 and has been so pervasive in common understandings of genetics that it has probably limited the perspectives of policymakers addressing genetic resources. If this is correct, then this article argues that founding a legislative, administrative and policy scheme on this misleading picture of bottom-up information flowing from DNA sequences may perpetuate perverse outcomes and (further)15 undermine the purpose and integrity of ABS schemes.

In addressing these matters, the article is structured as follows. The next part outlines the dimensions of the DSI issue in the CBD and Nagoya Protocol forums. The following part traces the developments of the use of informational language in genetics, detailing the current theoretical framework for information in philosophy and law and distinguishing between the ideals of classical and molecular genes so that simplistic and ultimately misleading conceptions of DNA sequence do not undermine the role and place of information in DNA sequences. The next part discusses the implications of these theoretical threads for the regulation of DSI in the context of ABS. The final part concludes that there already exists adequate potential in the current ABS scheme of CBD Contracting States and Nagoya Protocol Parties to implement ABS legislative, administrative and policy measures to regulate DSI as a genetic resource. Alternatively, those implementing existing national ABS can include terms and conditions as part of prior informed consent and mutually agreed terms addressing DSI (as some Contracting Parties have done already). While initiating such measures is possible, there are potential problems. The result will be a matrix of different laws, policies and practices among the CBD Contracting States and Nagoya Protocol Parties that will likely perpetuate perverse

6 Nagoya Protocol, Arts. 7 and 12.

7 See Young, “Drafting Successful ABS Contracts”; Humphries, “Survey of Access and Benefit-Sharing.”

8 See, for example, Gaffney, “Open Access to Genetic Sequence Data”; Williams, “Conservation Policy: Helping or Hindering”; Rourke,

“Policy Opportunities”; Laird, “Rethink the Expansion of ABS”; Lawson, “Information as the Latest Site”; Neumann, “Global Biodiversity Research”; Smith, “Biological Control”; Kupferschmidt, “Biologist Raise Alarm”; Bockmann, “Brazil’s Government Attacks”; Nijar,

“Implementation of the Nagoya ABS”; Prathapan, “When the Cure Kills”; Lawson, “The Future of Information”; Humphries, “A Tiered Approach.”

9 Lawson, “Information as the Latest Site,” 19–26.

10 CBD/DSI/AHTEG/2018/1/4, [25] and Annex ([1]).

11 Lawson, “The Future of Information,” 104.

12 See, for example, Colucci-D’amato, “End of the Central Dogma” (neurobiology); Maynard-Smith, “Concept of Information”

(developmental biology); Cooper, “Central Dogma of Cell Biology” (cell biology). See also Kay, “Who Wrote the Book of Life?”; Judson,

“The Eighth Day of Creation”; Kalmus, “Cybernetical Aspect.”

13 It broadly captures the explanation of biology and emphasises the minuteness of biological entities, although this is not an entirely clear episteme: see Kay, “The Molecular Vision of Life,” 4–6.

14 Sarkar, “Biological Information: A Skeptical Look,” 187. See also Falk, “Genetic Analysis: A History,” 175.

15 See Laird, “Rethink the Expansion of ABS”; Lawson, “Information as the Latest Site.”

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outcomes by controlling the potential uses of information and reducing the incentives for users of genetic resources to apply information in new and innovative ways. Consequently, this is likely to undermine the conservation and sustainable use of biodiversity because dealing with complicated laws, policies and practices will reduce the potential benefit-sharing opportunities for the uses of DSI. A preferable outcome would be an efficient and effective multilateral agreement balancing access and benefit-sharing that avoids the misleading picture of bottom-up information flowing from DNA sequences.

DSI as a CBD and Nagoya Protocol Issue

Formally recorded concerns about ABS and DSI in the CBD and Nagoya Protocol forums emerged in 201616 with the decision to establish an Ad Hoc Technical Expert Group on Digital Sequence Information on Genetic Resources (AHTEG-DSI).17 The AHTEG-DSI compiled and synthesised views about DSI and commissioned a fact-finding and scoping study to consider the concept and scope of DSI and how DSI was currently used.18 Significantly, the indicative and contextual information ‘that may be relevant to the utilization of genetic resources’ considered by the AHTEG-DSI included:

(a) the nucleic acid sequence reads and the associated data

(b) information on the sequence assembly, its annotation and genetic mapping. This information may describe whole genomes, individual genes or fragments thereof, barcodes, organelle genomes or single nucleotide polymorphisms

(c) information on gene expression

(d) data on macromolecules and cellular metabolites

(e) information on ecological relationships and abiotic factors of the environment (f) information on function, such as behavioural data

(g) structure, including morphological data and phenotype (h) information related to taxonomy

(i) modalities of use.19

The AHTEG-DSI concluded that more discussion about terminology was required to find a balance that could accommodate scientific, technological, market and other changes and provide legal certainty.20 Recognising a lack of consensus and common ground about the scope of the CBD and Nagoya Protocol and the likely consequences for DSI on benefit-sharing through technology transfer, partnerships and collaboration, information exchange and capacity development,21 the AHTEG-DSI continued together with an open-ended working group to develop modalities for sharing benefits from DSI22 and reported their findings in 2018.23 Reflecting the lack of agreement, the mandate of the AHTEG-DSI was extended, and they commissioned various additional studies on information traceability, sequence databases and domestic legal, administrative and policy ABS measures addressing DSI and benefit-sharing.24 The outcomes of the AHTEG-DSI were to be considered by the Open-Ended Intersessional Working Group to Support the Preparation of the Post-2020 Global Biodiversity Framework and at the next Conference of the Parties (COP) to the CBD and the Meeting of the Parties (MOP) to the Nagoya Protocol in 2020.25 However, the COP and MOP were postponed because of the global coronavirus pandemic.

The AHTEG-DSI has been a rich source of detail on DSI through its call for submissions and the commissioned peer-reviewed studies. The original commissioned scoping study identified the diversity of terms for DSI, including ‘resources in silico, genetic sequence data, genetic sequence information, digital sequence data, genetic information, dematerialized genetic resources, in silico utilization, information on nucleic acid sequences, nucleic acid information, and natural information

(emphasis in original).26 The study noted that more discussion of the terminology was required. It applied a basic conception of DSI as the order of nucleotides in a sequence that might be stored in a computer: ‘[DSI] is primarily the product of sequencing

16 CBD/COP/13/25, [321] and Decision XIII/16, [1]; CBD/NP/MOP/2/13, [153] and Decision 2/14, [1].

17 CBD/COP/13/25, [321] and Decision XIII/16, [4]. See also CBD/NP/MOP/2/13, [153] and Decision 2/14, [5].

18 CBD/DSI/AHTEG/2018/1/4, [17]. See also CBD/DSI/AHTEG/2018/1/2; CBD/DSI/AHTEG/2018/1/3.

19 CBD/DSI/AHTEG/2018/1/4, [23] and Annex ([2]).

20 CBD/DSI/AHTEG/2018/1/4, [23] and Annex ([12]).

21 A draft decision with entirely bracketed text: CBD/SBSTTA/22/12, [35] and Recommendation 22/1; CBD/DSI/AHTEG/2018/1/4, [23]

and Annex ([20]). See also Watanabe, “The Nagoya Protocol: The Conundrum”; Hammond, “Discussions on Sequence Information”; Scott,

“Workshop Report: Genetic,” 37–38.

22 CBD/SBSTTA/22/12, [18]–[19].

23 CBD/COP/14/14, [258] and Decision 14/20, [8 –[12]; CBD/NP/MOP/3/10, [162] and Decision 3/12.

24 CBD/COP/14/14, [258] and Decision 14/20, [11]. See also CBD/DSI/AHTEG/2020/1/7, [1].

25 CBD/COP/14/14, [258] and Decision 14/20, [12] and [237] and Decision 14/34, [2]. See also CBD/WG2020/3/4.

26 CBD/DSI/AHTEG/2018/1/3, 19–20.

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technologies that have become faster, cheaper, and more accurate in recent years. The aim of DNA sequencing is to determine the order in which each of the four DNA nucleotides is arranged in the molecule’.27

More recently, and extending previous work,28 another commissioned study considered the concept and scope of DSI and how DSI was currently used concluding that the proximity of the information to the underlying physical genetic resource provided a logical basis to group information that could comprise DSI as follows: ‘Group 1 – Narrow: DNA and RNA’; ‘Group 2 – Intermediate: (DNA and RNA) + proteins’; ‘Group 3 – Intermediate: (DNA, RNA and proteins) + metabolites’; and, ‘Group 4 – Broad: (DNA, RNA, protein, metabolites) + traditional knowledge, ecological interactions, [and so on]’.29 This study framed its discussion around the information flows represented by the ‘central dogma’ (DNA to RNA to protein to metabolites).30 The AHTEG-DSI also commissioned a combined study on databases and traceability of DSI31 that essentially limited their considerations to databases holding nucleotide sequence data and traceability in the core database infrastructure, the International Nucleotide Sequence Data Collaboration (INSDC).32 This included nucleic acid sequence reads (the sequence of nucleotides, such as CGAAAGACCGGC) and the associated data and information on the sequence assembly, annotation and genetic mapping.33 They found that some of these databases also included ‘subsidiary information’, which was broadly defined as information on gene expression, data on macromolecules and cellular metabolites, information on ecological relationships and other environmental data, functional data (e.g., behavioural data), structural data (e.g., morphological data and phenotype) and taxonomy data.34 The study concluded that the INSDC’s use of accession numbers (unique identifiers) facilitated database governance and traceability and that, at least in theory, this was a feasible mechanism for tracing nucleotide sequence data from a country or origin to a benefit-sharing user.35

The other AHTEG-DSI commissioned study on domestic measures addressing the commercial and non-commercial use of DSI and benefit-sharing found four kinds of legislative, administrative and policy measures: regulating DSI as a distinct object of ABS and separate from the physical genetic resources; regulating DSI as a part of the utilisation of physical genetic resources;

regulating DSI by requiring benefit-sharing (but not access) to cover the uses of DSI; and regulating DSI through other measures, such as compliance-related measures and monitoring mechanisms.36 The study found that some jurisdictions explicitly included ‘DSI’ language like ‘genetic information’, ‘genetic heritage’, ‘intangible components’, ‘gene sequences’,

‘sequence information’, ‘information’ and ‘information of genetic origin’, while others interpreted their existing ABS legislative, administrative and policy measures as including ‘DSI’, such as ‘genetic resources’, ‘genetic material’, ‘biological resources’, ‘associated knowledge’, ‘information of genetic origin’, ‘research results’ and ‘derivative’. However, the distinction between explicit and interpretive coverage was not necessarily clear.37 Where DSI was regulated as a distinct object of ABS and separate from the physical genetic resources, the ABS schemes extended broadly to include information associated with the genetic resources. For example, Malaysia’s ABS laws apply broadly to ‘biological resources’, which include ‘genetic resources, organisms, microorganisms, derivatives and parts of the genetic resources, organisms, microorganisms or derivatives’, ‘the populations and any other biotic component of an ecosystem with actual or potential use or value for humanity’ and ‘information relating to’ these ‘biological resources’. In addition, the definition of ‘derivative’ includes

‘information in relation to derivatives’.38 Kenya’s ABS laws apply to ‘access’, which means ‘obtaining, possessing and using genetic resources conserved, whether derived products and, where applicable, intangible components, for purposes of research, bio-prospecting, conservation, industrial application or commercial use’, where ‘intangible components’ are ‘any information held by persons that is associated with or regarding genetic resources’.39 Others regulate DSI by requiring benefit-sharing (but not access) obligations to cover the uses of DSI. For example, under India’s ABS laws, benefit-sharing obligations apply to

27 CBD/DSI/AHTEG/2018/1/3, 23.

28 CBD/COP/14/14, [258] and Decision 14/20, [11(a)].

29 CBD/DSI/AHTEG/2020/1/3, Annex (p. 32).

30 CBD/DSI/AHTEG/2020/1/3, Annex (pp. 10–12).

31 CBD/COP/14/14, [258] and Decision 14/20, [11(c)–(d)].

32 CBD/DSI/AHTEG/2020/1/4, Annex (p. 14). The report importantly notes that outside the INSDC databases, the ‘majority of such databases, after they are established during the project funding phase, are minimally, if at all, maintained, meaning webpages are infrequently updated, functions become defunct, or new data and bioinformatics tools are not added’ (p. 18).

33 CBD/DSI/AHTEG/2020/1/4, Annex (p. 14).

34 CBD/DSI/AHTEG/2020/1/4, Annex (p. 14).

35 CBD/DSI/AHTEG/2020/1/4, 64–66.

36 CBD/DSI/AHTEG/2020/1/5, Annex (pp. 9–11).

37 CBD/DSI/AHTEG/2020/1/5, Annex (pp. 11–12).

38 Access to Biological Resources and Benefit-Sharing Act 2017 (Malaysia), s. 4.

39Environmental Management and Co-ordination (Conservation on Biological Diversity and Resources, Access to Genetic Resources and Benefit Sharing) Regulations 2006 (Kenya), reg. 2.

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‘biological resource occurring in India or knowledge associated thereto’ for ‘research or for commercial utilisation or for bio- survey and bio-utilisation’,40 where ‘research’ means the ‘study or systematic investigation of any biological resource or technological application, that uses biological systems, living organisms or derivatives thereof to make or modify products or processes for any use’; ‘commercial utilisation’ means ‘end uses of biological resources for commercial utilisation such as drugs, industrial enzymes, food flavours, fragrance, cosmetics, emulsifiers, oleoresins, colours, extracts and genes used for improving crops and livestock through genetic intervention, but does not include conventional breeding or traditional practices in use in any agriculture, horticulture, poultry, dairy farming, animal husbandry or bee keeping’; and ‘bio-survey and bio- utilisation’ means the ‘survey or collection of species, subspecies, genes, components and extracts of biological resource for any purpose and includes characterisation, inventorisation and bioassay’.41 The Indian law also provides that ‘[n]o person shall, without the previous approval of the National Biodiversity Authority, transfer the results of any research relating to any biological resources occurring in, or obtained from, India’.42 While there is no consensus apparent in the existing practices about the best ways to regulate DSI, there are, as the examples demonstrate, various forms of genetic information already subject to regulation in implementing CBD and Nagoya Protocol-consistent ABS schemes.

In addition to these commissioned studies, submissions of views and information to clarify the concept of DSI and benefit- sharing arrangements from using DSI have been made by CBD Contracting Parties and others, including other governments, Indigenous Peoples and local communities, relevant organisations and stakeholders.43 A range of responses have been submitted essentially in three groupings: those arguing that DSI should not be a part of the CBD and Nagoya Protocol;44 those favouring some accommodation of DSI;45 and those favouring or already including DSI in their legal, policy and administrative ABS arrangements.46 As a broad generalisation, technologically rich CBD Contracting Parties favour DSI not being a part of the CBD and Nagoya Protocol, and technologically poor CBD Contracting Parties favour DSI being accommodated or included in the CBD and Nagoya Protocol. As a useful summary of the way forward, the recent Open-Ended Working on the Post-2020 Global Biodiversity Framework considered a typology of possible regulatory options (although traditional knowledge associated with genetic resources was not addressed):47 ‘Option 0: Status Quo’, addressing DSI under the existing arrangements through domestic ABS laws, policies and processes; ‘Option 1: DSI Fully Integrated into the [CBD] and the Nagoya Protocol’, addressing DSI as a genetic resource under the CBD and Nagoya Protocol and as an obligation under those agreements and implemented in domestic ABS laws, policies and processes; ‘Option 2: Standard [Mutually Agreed Terms]’, addressing DSI through an obligation to share benefits from the uses of DSI without restricting access to DSI itself through some kind of agreement with standard terms and conditions; ‘Option 3: No [Prior Informed Consent], No [Material Transfer Agreement]’, addressing DSI by requiring a payment or contribution for access or use of the DSI into a multilateral fund without the need for prior informed consent or mutually agreed terms and ABS contracts; ‘Option 4: Enhanced Technical and Scientific Cooperation’, democratising access and use of DSI so that each country has the capacity and opportunity to access and use DSI; and ‘Option 5: No Benefit Sharing from DSI’, no mechanisms are proposed and there is no benefit-sharing from the use of DSI.48

Returning to the AHTEG-DSI and the indicative and contextual information ‘that may be relevant to the utilization of genetic resources’,49 the commissioned study essentially considered four kinds of information according to ‘the flow of information from a genetic resource, particularly the degree of biological processing and proximity to the underlying genetic resource, to provide a logical basis to group information that may comprise DSI’.50 Underpinning this ‘logical basis to group information’

was a particular conception of information in genetics founded in an ideal of the ‘central dogma’,51 which expresses genetic information as ‘nucleotide sequence information associated with transcription’, ‘protein sequence’ information associated with translation, ‘information associated with transcription and translation’ and ‘metabolites and biochemical pathways, thus comprising information associated with transcription, translation and biosynthesis’ and ‘extends to behavioural data, information on ecological relationships and traditional knowledge, thus comprising information associated with transcription,

40 Biological Diversity Act 2002 (India), s. 3.

41 Biological Diversity Act 2002 (India), s. 2.

42 Biological Diversity Act 2002 (India), s. 4.

43 CBD/COP/14/14, [258] and Decision 14/20, [9].

44 Examples include Australia, Canada, Japan and Korea.

45 Examples include the European Union and its Member States and Switzerland.

46 Examples include the African Union Commission on behalf of the African Group, Brazil, Ethiopia, India, Iran, Madagascar and South Africa.

47 CBD/WG2020/3/4, Annex II (p. 13).

48 CBD/WG2020/3/4, Annex II (pp. 15–17).

49 CBD/DSI/AHTEG/2018/1/4, [23] and Annex ([2]).

50 CBD/DSI/AHTEG/2020/1/3, Annex (p. 32). See also CBD/WG2020/3/4, Annex I (p. 8).

51 CBD/DSI/AHTEG/2020/1/3, 10–15.

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translation and biosynthesis, as well as downstream subsidiary information concerning interactions with other genetic resources and the environment as well as its utilization, among other subsidiary information’.52 Importantly, the study also addressed the

‘degree of biological processing and proximity to the underlying genetic resource’ to distinguish between ‘data’ and

‘information’, the latter information being processed data.53 The issue of the broader concept of biological information is addressed next. However, first, it is important to make a distinction here between information about DNA sequences and information in DNA sequences.

Information about DNA sequences is the vast quantity of information produced, collected, stored, accessed, managed and manipulated, including the order of nucleotides in a sequence, how the sequencing was conducted, annotations and functional analysis. This information is the subject matter of the information sciences bioinformatics applied to genetics:

What makes biology an information science in this sense is not anything about the nature of genes, but the fact that contemporary biology works with vast bodies of data that the unaided human mind is incapable of processing effectively.54

The CBD already has an extensive mechanism to address this information about DNA sequences that is independent of the ABS obligations.55 Essentially, the CBD has a general obligation to promote the exchange of information on the ‘results of technical, scientific and socio-economic research’, ‘training and surveying programmes’, ‘specialized knowledge’ and

‘[I]ndigenous and traditional knowledge as such and in combination with the technologies [‘relevant to the conservation and sustainable use of biological diversity or make use of genetic resources’]’56 and ‘where feasible, include repatriation of information’.57 There is a clearing house mechanism ‘to promote and facilitate technical and scientific cooperation’58 realised through decentralised databases and websites (information hubs) and national government websites.59 The Nagoya Protocol Access and Benefit Sharing Clearing House is a part of the CBD’s clearing house mechanism and applies only to ABS arrangements and ‘access to information made available by each Party relevant to the implementation of this [Nagoya]

Protocol’.60 The CBD’s Clearing House Mechanism (including the Nagoya Protocol Access and Benefit Sharing Clearing House), linked sites and sites linked to those sites set out information about DNA sequences and genetic resources more broadly.61

In contrast, information in DNA sequences is ‘a theoretical entity which exists in the genome and explains biological phenomena’.62 Information in DNA sequences is the information in genetic resources as opposed to the information about genetic resources.

DNA Sequences as Information

It is uncontroversial that DNA is a linear sequence of molecules that can be presented as syntactic information in the language of the genetic code.63 Letters of the alphabet making words that are joined into sentences, paragraphs and chapters represent syntactic information in language (here English). Similarly, photographs, music and computer programs are all, or can be rendered into, linear sequences of syntactic information in language (i.e., 0s and 1s of binary code). The proposition here is that because linear sequences in the form of words in sentences, paragraphs and chapters (and also photographs, music and computer programs) are information, then, similarly, DNA molecules in a linear sequence with a code represent information.64 The question then is whether DNA molecules can actually be information.

52 CBD/DSI/AHTEG/2020/1/3, 32. See also CBD/WG2020/3/4, Annex I (p. 8).

53 CBD/DSI/AHTEG/2020/1/3, 42–43.

54 Griffiths, “Genetics and Philosophy,” 145.

55 For an overview, see Lawson, “Information as the Latest Site,” 19–26.

56 CBD, Arts. 16.1 and 17.1.

57 CBD, Art. 17.2.

58 CBD, Art. 18.3.

59 See UNEP/CBD/WGRI/5/3/Add.2; UNEP/CBD/COP/12/11.

60 Nagoya Protocol, Art. 14. UNEP/CBD/COP/10/27, [103] and Annex (Art. 14(1); Decision X/1, pp. 85–109). See also UNEP/CBD/COP/12/6, [51]–[58]; UNEP/CBD/ICNP/3/6.

61 See, for example, the Australian Government’s Department of Environment and Energy website that sets out information about national strategies for biodiversity conservation, regulation and links to other websites that hold research and publications about research that includes information about genetic resources: http://www.environment.gov.au/biodiversity.

62 Griffiths, “Genetics and Philosophy,” 144–145.

63 See Crick, “Central Dogma of Molecular Biology”; Crick, “On Protein Synthesis.”

64 See, for example, Barbieri, “Definitions of Information and Meaning”; Barbieri, “Life and Semiosis.”

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The ideal of a DNA sequence as information traces back to at least 195365 and the ‘central dogma’ that information flows from DNA to RNA to proteins but not the other way out of proteins.66 According to this account, the organism’s genome accumulates information (through the mechanisms of evolution) for transmission to the next generations.67 The organism itself is merely the reservoir and transmitter of information.68 Taken literally, a DNA sequence as information means that the arrangements of As, Ts, Gs and Cs represent the raw data, and they are themselves information. This is consistent with the ideal of life as information and programmable Boolean switches.69 This might appear intuitively correct given the explosion of bioinformatics as a technological discipline exploiting information.70 Unfortunately, this notion overlooks the complexity of genetics and gives undue weight to a particular conception of genotype as the causative (or purposeful) explanation (as addressed in detail below).

The trajectory of this debate is important because developing regulatory schemes based on a particular perspective or preference regarding an unsettled theoretical foundation is likely to lead to bad laws and unforeseen consequences. At the heart of this problem is finding a common understanding for the term ‘information’ in genetics and how this might be addressed by law because, very crudely, it is not clear whether ‘information’ is addressing these molecules literally or metaphorically.71

A good entry point into this ongoing debate, and an obviously very brief account,72 is Charles Darwin’s 1859 theory of natural selection that set aside the idea that species were immutable, static and designed by a god. Darwin instead introduced the idea that species had adapted to their environments over many generations.73 While Darwin posited that evolution and inheritance were linked, he also accepted that he was unable to explain the mechanism by which traits were inherited.74 In 1865, Gregor Mendel provided such an account, positing from his experiments with peas that dominant and recessive elements were inherited.

He traced those elements through hybrids as different constitutions and groupings of elements (‘Faktoren’).75 Mendel’s insight was to mediate the relationship between genotypes (‘genes’) and their phenotypes (‘unit characters’)76 by assigning the unobservable genotype to a phenotype (i.e., traits, such as seed texture, seed colour, pollen texture and pollen colour) and tracking the ‘dominant’ and ‘recessive’ phenotypes across hybridising crosses.77 In this sense, Mendel’s elements were necessary for his explanation to work,78 and as such, the Mendelian gene (albeit Mendel was no Mendelian)79 refers to the unit of inheritance (‘Zellelemente’)80 that predicted the apparent characters across generations. At its most simple, the Mendelian gene is an account of a mechanism for observed phenotypes from sexual crosses. This account posits a ‘gene’ (coined by Wilhelm Johannsen)81 to be an undefined unit of inheritance transmitted across generations and links the phenotype to a genotype.82 Here the genotype was the speculated and inferred cause of the observed phenotypes, with no explanation of the material and instrumental manifestation of the Mendelian gene itself.83 Importantly, however, ‘what were studied were character differences, not characters, and what explained them were differences in genes, not the genes themselves’.84 As a unit of inheritance, the Mendelian gene was a theoretical explanation of two kinds: first, the heritable factors an offspring receives, one from each parent in sexual crosses, albeit not an observable entity but an explanation of the observations of segregation

65 Watson, “Molecular Structure of Nucleic Acids,” 737: ‘It has not escaped our notice that the specific pairing we have postulated immediately suggests a possible copying mechanism for the genetic material.’

66 Crick, “Central Dogma of Molecular Biology”; Crick, “On Protein Synthesis.”

67 Maynard-Smith, “Concept of Information,” 185–192.

68 Morange, “History of Molecular Biology,” 2.

69 For an elegant account of this perspective, see Rosenberg, “Darwinian Reductionism,” 56–93.

70 See, for example, Ranganathan, Encyclopedia of Bioinformatics.

71 Griffiths, “Genetics and Philosophy,” 146–147.

72 The history of genetics is well traversed. See, for example, Falk, “Genetic Analysis: A History”; Carlson, “Mendel's Legacy”; Keller, “The Century of the Gene”; Morange, “History of Molecular Biology”.

73 Darwin, “On the Origin of Species.” For an overview, see Bowler, “Evolution: The History of an Idea.”

74 Darwin, “On the Origin of Species,” 19–20.

75 Mendel, “Versuche über Pflanzenhybriden,” 42.

76 For language developed by Wilhelm Johannsen, see Johannsen, “The Genotype Conception of Heredity.” See also Roll-Hansen, “Sources of Wilhelm Johannsen’s Genotype Theory.”

77 This is the passing over of a more complicated and intriguing moment in the history of genetics: see Falk, “Mendel’s Impact.”

78 Griffiths and Stotz, “Genetics and Philosophy,” 15.

79 Olby, “Mendel no Mendelian?”

80 Mendel, “Versuche über Pflanzenhybriden,” 42.

81 See Johannsen, “Elemente der exakten Ereblichkeitslehre.”

82 See Wanscher, “Analysis of Wilhelm Johannsen’s.”

83 For an account of this perspective, see Waters, “Genes Made Molecular,” 169–174 and the references therein.

84 Waters, “Genes Made Molecular,” 172.

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and independent assortment (the classical gene); and second, the material and instrumental entity of heredity (the molecular gene).85

Both forms of the Mendelian gene persist,86 although modern genetic practitioners often conflate the two forms:

When molecular biologists focus on nucleotide sequences, they think of genes in molecular concept. But at earlier stages of investigation, when they have not gotten close to specifying nucleotide sequences, they tend to think of genes in terms of the rougher-grained classical concept.87

Rather than thinking of classical genes and molecular genes as separate theories,88 most genetics practitioners consider a continuous theory addressed at two levels of resolution, with the classical genes being an ‘organic extension’ of the molecular gene.89 Conceived this way, genetics is a reductionist, bottom-up account based in a physical sciences methodology using numerical analyses, with the outcome that the methodological and conceptual understanding of heredity is reduced to the sum of the physical and chemical properties of its building blocks, and that is DNA as the molecular gene, so the genotypes are the causes of phenotypes.90 For the present purposes, it is significant that the later biochemical and molecular biological account of the material and instrumental manifestation of the gene overlooked this classical account of the units of a genetic and non- genetic context that resulted in the observed phenotype.91 Put simply, ‘molecular biologists can now determine the exact molecular identity of the relevant differences and explain how in general such differences produce phenotypic difference within a genetic context’ (emphasis added).92 This becomes clear when tracing the ideal of the material and instrumental manifestation of the molecular gene as opposed to the classical gene. This is important because the molecular gene has taken precedence as the account of genetics and gained a popular appeal93 that overlooks much of the intriguing complexity and the role of other non-genetic (epigenetic)94 factors in the observed phenotype, such as environmental effects.

The key moments in tracing the primacy of the material and instrumental manifestation of the molecular gene might be, hopefully uncontroversially,95 stated as: Wilhelm Hofmeister first observed chromosomes in the 1840s;96 Wilhelm Roux speculated that chromosomes are the carriers of inheritance in the 1880s;97 Walter Sutton and Theodor Boveri correlated the action of chromosomes with the apparent results of Mendel;98 Wilhelm Johannsen used the word ‘gene’ in 1909 to describe the fundamental physical and functional unit of heredity;99 in 1910, Thomas Morgan proposed that genes (then called ‘factors’) are located on specific chromosomes;100 Alfred Sturtevant provided a linkage map of genes in 1913;101 Theophilus Painter produced a cytological mapping of fruit fly salivary glands to localise genes at specific chromosome locations in 1934;102 George Beadle and Edward Tatum proposed the ‘one gene’ hypothesis based on their experiments showing that specific steps in metabolic

85 For an engaging discussion of these different uses of the gene concept leading to Gregor Mendel being characterised as a methodological reductionist, the later re-discovery of Mendel’s work and Hugo de Vries applying the concept to the material and causal elements as a conceptual reductionist, see Falk, “Genetic Analysis: A History,” 4. See also Kitcher, “1953 and all That,” 336.

86 See Carlson, “Mendel's Legacy.”

87 Waters, “Genes Made Molecular,” 183.

88 See, for example, the different classical phenotypic marker gene-P and the molecular sequence gene-D: Moss, “What Genes Can’t.” See also Kitcher, “1953 and all That,” 336–337.

89 Falk, “Genetic Analysis: A History,” 3–4.

90 See Falk, “Mendel’s Impact,” 216–226. See also Fuerst, “The Role of Reductionism.”

91 See Falk, “Mendel’s Impact,” 229–233.

92 Waters, “Genes Made Molecular,” 183–184.

93 See, for an example of a simplistic (and incorrect) representation of information flows: Wright, “DNA → RNA.”

94 As used here and might be used in discussing molecular biology, ‘epigenetic’ means ‘the study of changes in gene expression t hat are mitotically heritable (via somatic cells) or meiotically heritable (via germ cells), and that do not entail changes in DNA sequence’: Griffiths,

“Genetics and Philosophy,” 113.

95 There are various accounts. See, for example, Portin, “The Evolving Definition of the Term Gene”; Allen, “Naturalists and Experimentalists”; Carlson, “The Gene: A Critical History.”

96 Hofmeister, “Ueber die Entwicklung des Pollens”; Witty, “Pollen Development.” See also Kaplan, “The Genius of Wilhelm Hofmeister,”

1650–1652.

97 Roux, “Ueber die Bedeutung der Kerntheilungsfiguren.” See also Hamburger, “Wilhelm Roux: Visionary,” 232–233.

98 Sutton, “The Chromosomes in Heredity”; Boveri, “Über Mehrpolige Mitosen.” Noting that this claim is contested, see Martins, “Sutton and Boveri.”

99 Johannsen, “Elemente der exakten Ereblichkeitslehre.” See also Johannsen, “The Genotype Conception of Heredity.”

100 Morgan, “Sex-limited Inheritance,” 120–121. This proposition was more fully articulated in Morgan, “Mechanism of Mendelian Heredity.” Note, however, that Morgan appears to have been influenced by others: see Edwards, “Robert Heath Lock” (proposing that genes lying on a chromosome might account for linkages).

101 Sturtevant, “The Linear Arrangement.”

102 Painter, “A New Method,” 175.

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pathways were disrupted by mutations in 1941;103 Oswald Avery demonstrated that DNA was the hereditary material causing the heritable changes in 1944;104 Erwin Chargaff clarified that the number of guanine and cytosine units and the number of adenine and thymine units were the same, hinting at the base pair makeup of the DNA;105 James Watson and Francis Crick proposed the double helix structure for DNA in 1953;106 Seymour Benzer proposed the conception of genes as linear structures along chromosomes (rather than being like beads on a necklace, they are instead divisible into smaller units of mutation and recombination) in 1955;107 and, the elucidation of the genetic code by Crick and others in 1961.108 The end of this track is the ideal of a linear molecular gene where the DNA sequence is considered the genotype and causative agent for the observed phenotype.

Intriguingly, Watson and Crick, in proposing the double helix structure for DNA in 1953, speculated about the ‘possible copying mechanism for the genetic material’109 and ‘the precise sequence of the bases in the code that carries the genetic information’.110 This was essentially entrenching using information language in molecular biology that had started with terms such as ‘words’,

‘codes’, ‘messages’ and ‘texts’ in the 1930s and took solid hold in the 1940s.111 Crick’s speculation later matured to the generalised rule for the informational transfer from one polymer to another (DNA to RNA, RNA to DNA, RNA to RNA, DNA to DNA and RNA to protein but not protein to protein,112 protein to DNA and protein to RNA)113 so that ‘once “information”

has passed into protein it cannot get out again’, where ‘information’ means ‘the precise determination of sequence, either of bases in the nucleic acid or of amino acid residues in the protein’ (emphasis in original).114 There are two parts to this claim that played out over the following decades. First, there is a coded sequence specificity between the DNA and the transcribed RNA and the translated polypeptide (sequence hypothesis).115 Second, the expression of the DNA sequence determines the RNA or a protein product such that all products are informed (specified or caused) by the DNA sequences (central dogma).116 Some of the details here matter: Crick accepted that protein synthesis involved ‘the flow of energy, the flow of matter, and the flow of information’, and his focus was the ‘information’.117 Crick later stated that:

it was abundantly clear by that time that a protein had a well-defined three-dimensional structure, and that its activity depended crucially on this structure, it was necessary to put the folding-up process on one side, and postulate that, by and large, the polypeptide chain folded itself up.[118] This temporarily reduced the central problem from a three dimensional one to a one dimensional one … The principal problem could then be stated as the formulation of the general rules for information transfer from one polymer with a defined alphabet to another.119

The key advances reinforcing these determinist and informational explanations of the now linear molecular gene, again hopefully uncontroversially, were: François Jacobs, Jacques Monod, Sydney Brenner, François Gros, Francis Crick and others’

103 Beadle, “Genetic Control of Biochemical Reactions,” 505–506.

104 Avery, “Studies on the Chemical Nature,” 149–150. See also Wyatt, “When Does Information Become Knowledge”; Hershey,

“Independent Functions of Viral Protein.”

105 Chargaff, “Chemical Specificity of Nucleic Acids.”

106 Watson, “Molecular Structure of Nucleic Acids,” 737.

107 Benzer, “Fine Structure of a Genetic Region,” 347.

108 See Crick, “General Nature of the Genetic Code.”

109 Watson, “Molecular Structure of Nucleic Acids,” 737.

110 Watson, “Genetical Implications,” 965.

111 See Kay, “The Molecular Vision of Life,” 3–57.

112 Crick did admit some ambiguity here, presciently considering prions: ‘There is, for example, the problem of the chemical nature of the agent of the disease scrapie’: Crick, “Central Dogma of Molecular Biology,” 562. Also note that there is evidence of proteins catalysing amino acid polymerisation: see, for example, Rout, “Prebiotic Template-directed Peptide Synthesis.”

113 Note that Crick asserts that reverse transcriptase using viral RNA as a template for DNA synthesis was not a reversal of the central dogma but, rather, a ‘misunderstanding’: Crick, “Central Dogma of Molecular Biology,” 561. See also Morange, “What History Tells Us.”

114 Crick, “On Protein Synthesis,” 153. See also Crick, “Central Dogma of Molecular Biology,” 562. See also Sarabhai, “Co-linearity of the Gene.”

115 ‘The Sequence Hypothesis ... in its simplest form assumes that the piece of nucleic acid is expressed solely by the sequence of its bases, sequence is a (simple) code for the amino acid sequence of a particular:’ Crick, “On Protein Synthesis,” 152.

116 ‘The Central Dogma of molecular biology deals with the detailed residue-by-residue transfer of sequential information. It states that such information cannot be transferred from protein to either protein or nucleic acid’: Crick, “Central Dogma of Molecular Biology,” 561. For analyses of the various interpretations, see Camacho, “Central Dogma is Empirically Inadequate.” See also Thieffry, “Forty Years under the Central Dogma.”

117 Crick, “On Protein Synthesis,” 139–140.

118 This was later shown to be correct because the polypeptide chain folded itself into the thermodynamically most stable form. See Anfinsen,

“The Kinetics of Formation.”

119 Crick, “Central Dogma of Molecular Biology,” 561.

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discovery of mRNA in 1960;120 the genetic code linking triplets of nucleotides (codons) to specific amino acids in 1961;121 Jacobs and Monod’s explanation of a regulation mechanism (the lac operon) on a linear DNA molecule accounting for the relationship between DNA, RNA and proteins and pointing to a hierarchical network of regulation in 1962,122 Jim Shapiro and others’ isolation of a bacterial gene (the lac operon) in 1969,123 Howard Temin and David Baltimore’s discovery of the enzyme that reversed transcription process making DNA from an RNA template in 1970,124 David Jackson, Robert Symons and Paul Berg making recombinant DNA molecules in 1972,125 Herbert Boyer and Stanley Cohen showing that engineered DNA molecules could be cloned in foreign cells in 1973,126 and, then the sequencing of various genomes, including the bacteriophage φX174 in 1978,127 Haemophilus influenzae in 1995,128 yeast Saccharomyces cerevisiae in 1996,129 the nematode Caenorhabditis elegans in 1998,130 the fruit fly Drosophila melanogaster in 1999131 and the human genome in 2000.132 The end of this track of enquiry was to cement the ideal of a linear molecular gene where the DNA sequence was popularly considered the genotype and causative agent for all the observed phenotypes.

The appeal of this approach was the focus on the simple explanatory power133 of the information comprised by DNA both as a store of evolutionary accumulated changes and as a master plan for cell development and performance.134 This was also the logic of a reductive physics and chemistry account where the molecule is the semantics of the transmitted information: ‘the polypeptide phenotype is determined by the polynucleotide genotype’.135 This has been, as the tracing of key advances above illustrates, amazingly heuristically successful. The key point, however, is that framed this way, the DNA sequence is conceived as a repository of meaning, aboutness and content (intentional or semantic information) regarding complex phenotypes so that the genotypes are privileged in the causes of phenotypes and the ideal that there is information in the DNA sequence.136 The problem remains, however: If there is information in the DNA sequence, what kind of information is it?

Information in DNA Sequences

Even when Watson and Crick were proposing their DNA structure, the information metaphors were known to be problematic.137 The postgenomic era following the release of the draft human genome sequence in 2001138 revealed that the relationship between the 20,000–25,000 structural genes and the approximately 1,000,000 polypeptides of the proteome139 was a lot more complicated than a mere linear sequence of nucleotides corresponding to a linear order of the gene products (RNA and amino acids in a polypeptide). Since then, the disruption of information flows has been repeatedly demonstrated, revealing that the DNA sequence and other factors (including non-genetic factors) are the contributors to RNA and protein sequence specificity (both quality and quantity): genes comprising complex regulatory networks affected by non-genetic factors result in a range of stable and robust phenotypes;140 epigenetic acetylation, phosphorylation and methylation markings can regulate gene transcription;141 a range of cis- and trans-acting factors interact with the linear DNA sequence (e.g., transcription factors, promotors, activators, repressors, enhancers, silencers, and splicing factors), resulting in a diversity of RNAs (e.g., mRNA, rRNA, tRNA, lncRNA and RNAi) and protein forms/splice variants;142 reordering of the linear DNA sequences through

120 See Morange, “History of Molecular Biology,” 139–149 and the references therein.

121 Crick, “General Nature of the Genetic Code.” See also Crick, “The Genetic Code – Yesterday.”

122 Jacobs and Monod, “Genetic Regulatory Mechanisms.”

123 Shapiro, “Isolation of Pure lac Operon.”

124 Baltimore, “RNA-dependent DNA Polymerase”; Temin and Mizutani, “RNA-dependent DNA Polymerase.”

125 Jackson, “Biochemical Method for Inserting New Genetic Information.”

126 Cohen, “Construction of Biologically Functional Bacterial Plasmids.”

127 Sanger, ‘Nucleotide Sequence of Bacteriophage φX174.”

128 Fleischmann, “Whole-Genome Sequencing.”

129 See Goffeau, “Life with 6000 Genes.”

130 The C. elegans Sequencing Consortium, “Genome Sequence of the Nematode C. elegans.”

131 Adams, “The Genome Sequence of Drosophila melanogaster.”

132 Venter, “The Sequence of the Human Genome”; Human Genome Sequencing Consortium, “Initial Sequencing and Analysis of the Human Genome.”

133 Compellingly ‘simple and pedagogic’ power that made molecular biology attractive: Morange, “History of Molecular Biology,” 174.

134 Maynard-Smith, “Concept of Information,” 178–181. See also Schaffner, “The Watson-Crick Model.”

135 Falk, “Genetic Analysis: A History,” 260.

136 See Griffiths, “The Fearless Vampire Conservator,” 175–198.

137 See Keller, “Refiguring Life,” 10; Kay, “Who Wrote the Book of Life?” 621–622. See also Fogle, “Information Metaphors.”

138 Venter, “The Sequence of the Human Genome.”

139 Muller, “Annotating the Human Proteome,” 176.

140 See, for example, Wagner, “The Role of Robustness in Phenotypic Adaptation.”

141 See, for example, Cavalli, “Advances in Epigenetics.”

142 See, for example, Morris, “The Rise of Regulatory RNA.”

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frameshifting, programme slippage or bypassing and codon redefinition;143 and RNA editing resulting in a significantly larger transcriptome.144 These examples confirm that there is not necessarily a consistent nexus between the DNA sequence as the sole source of information flowing from the DNA to an RNA and a protein.145 Put simply, the evidence now clearly shows that there is not always a direct correspondence between the DNA sequence and protein because there is processing and modification of that sequence on the way from DNA to protein,146 and modification of that processing and function can be from outside the DNA sequences (and particularly from the environment).147 This calls for a nuanced depiction of DNA as information because a DNA sequence alone (genotype) cannot cause the whole organism. There are other contributing causes, such as the environment, and thus, the classical gene (observed phenotype) is not necessarily only caused by the molecular gene (deterministic and reduced to a DNA sequence). There are potentially better accounts of the information in DNA sequences, as argued in the following paragraphs.

A useful starting point is Shannon information theory, which posits a simple quantitative framework for describing correlations:

two variables are correlated in some sense where the output of the channel depends on the input.148 Here the correlation termed information has a special and technical mathematical sense that treats both sense and non-sense input messages as the same.149 Taking this a step further, the information might be considered to have some content in the sense of natural signs and indicators150 and that there is a correlation between a DNA sequence and an observed phenotype151 with information conveyed in the sense of its natural meaning.152 Thus:

The key currency in information theory is the entropy H(X) of a random variable X. The entropy is a measure of uncertainty in the realization of X. If X takes on value Xi with probability pi, the entropy H(X) = Σipilog pi. The key statistic in information theory is the mutual information I(X;Y) between two random variables X and Y. The mutual information, defined as I(X;Y) = H(X)-H(X|Y), measures how much we learn about the value of X by knowing Y.153

If the DNA sequence is information—in this sense of information as the correspondence between the linear sequences of DNA nucleotides that specifies the linear sequences of RNA ribonucleotides and the linear sequences of amino acids (protein)—then Shannon information theory has some application with the input DNA sequence, the output amino acid sequence and the information transfer from DNA to amino acid modelled as the communication channel.154 This accepts that DNA sequences do have a limited causal role155 but not a broader intentional or semantic role.156 Using Crick’s words, this is ‘the general rules for information transfer from one polymer with a defined alphabet to another’,157 where ‘information was “merely a convenient shorthand for the underlying causal effect”, namely the “precise determination of sequence”’.158 This may also be conceived as a causal specificity between the DNA, the RNA and the coded amino acids,159 representation of the RNA and amino acids in the DNA sequence and the correlation information between the DNA sequence, RNA and amino acids160—the ‘information that specifies the product is no longer carried in a three-dimensional structure but instead by the linear, one-dimensional order

143 See, for example, Schmucker, “Dscam and DSCAM.”

144 See, for example, Tang, “Biological Significance of RNA Editing.”

145 Stotz, “With ‘Genes’ Like That,” 905; Stotz, “Molecular Epigenesis,” 542. For a different view see, for example, Baetu, “A De fense of Syntax,” 713–714; Waters, “Genes Made Molecular,” 174–182.

146 See Falk, “Genetic Analysis: A History,” 262.

147 See Griffiths, “Genetic Information,” 395–396.

148 See Shannon, “Mathematical Theory of Communication.” See also Fabris, ‘Shannon Information Theory.”

149 Shannon, “Mathematical Theory of Communication,” 99.

150 Dretske, “Epistemology and Information,” 31.

151 See Godfrey-Smith, “Information in Biology,” 104. Noting, however, that a sign, an object and an interpretant may actually be required for there to be any meaningful correlation. Hence, ‘while tree rings might be a source of quantitative information, they would not mean anything about a tree’s age unless there were an agent present who understood how tree rings are produced and how they relate to yearly growth’: Kumar, “Information, Meaning, and Error,” 91.

152 Grice, “Meaning.”

153 Bergstrom, “The Transmission Sense of Information,” 161.

154 See, for example, Bynum, “Informational Metaphysics,” 205; Román-Roldán, “Application of Information Theory to DNA Sequence Analysis,” 1188. Noting, of course, this might all be a misunderstanding: see Ben-Naim, “Entropy and Information Theory,” 1185–1189.

155 See, for example, Godfrey-Smith, “Information, Arbitrariness, and Selection,” 204. For a contrary view, see Kitcher, “Battling the Undead”

(arguing genetic coding has no explanatory weigh).

156 For other views, see Rosenberg, “Is Epigenetic Inheritance a Counterexample”; Fantini, “Of Arrows and Flows”; Weber, “The Central Dogma as a Thesis.”

157 Crick, “Central Dogma of Molecular Biology,” 561.

158 Stotz, “Biological Information, Causality and Specificity,” 369. See also Griffiths, “Genetic, Epigenetic and Exogenetic Information,” 2.

159 See Griffiths, “Measuring Causal Specificity”; Pocheville, “Comparing Causes,” 93–102.

160 See Shea, “Representation in the Genome,” 314.

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