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Paper Citation Record · LEDGER

Large Language Model is Secretly a Protein Sequence Optimizer

As of 20 August 2026, this Paper Citation Record lists 27 of 27 outbound references and 3 inbound Pith citation observations for arXiv:2501.09274.

A citation records a reference. It does not transfer a finding from one paper to another.

pith.paper-citation-record.v1
2501.09274 v2

Coverage vector

measured 27 of 27 reference resolution

Typed states for the displayed outbound observations.

Source: paper_references, paper_reference_links, observed 2026-08-10T20:10:52.893429Z

measured 30 of 30 standing notices

One-hop event checks from named stored sources.

Source: scholarly_work_events, retraction_status_cache, observed 2026-08-20T06:33:59.587034+00:00

measured 3 of 3 inbound itemization

Pith citing papers itemized under the disclosed page cap.

Source: paper_references, paper_reference_links, observed 2026-08-04T20:55:41.916450Z

measured 0 of 1 external citation measurements

A source-named dated measurement, never combined with another source.

Source: arxiv_reference, observed 2026-07-03T04:07:36.964600Z

Reference resolution

27 of 27 outbound references displayed

  • verified exact1
  • verified fuzzy14
  • unresolved12
  • parse uncertain0
  • malformed identifier0
  • metadata mismatch0

External citation measurements

No source-named external measurement is stored.

Outbound references

Observation b4cb0224-a333-4f9c-b4e9-fb17880c889d · outbound

This paper cites Design by directed evolution.

Large Language Model is Secretly a Protein Sequence Optimizer Design by directed evolution

Reference 1

Resolution
unresolved
no resolver link, observed 2026-08-10T20:10:52.799942Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.799942Z digest=sha256:4a4b65dff119fa0e66c3d90bc475d9bb17a25916deaf8d695f821f5191798598

Observation 11f9d901-d3f7-4c61-a4f9-bac6fd0be97e · outbound

This paper cites Conditioning by adaptive sampling for robust design.

Large Language Model is Secretly a Protein Sequence Optimizer Conditioning by adaptive sampling for robust design

Reference 2

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.258807Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.803499Z digest=sha256:c90ea1c72fc9f557932977c0f1f5bd2c9c50f9233fa722b10be5d1fd5f49b367

Observation e0016b4e-82e3-40bd-bf3c-f7dabb0cbd71 · outbound

This paper cites Deep diversification of an aav capsid protein by machine learning.

Large Language Model is Secretly a Protein Sequence Optimizer Deep diversification of an aav capsid protein by machine learning

Reference 3

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.247851Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.806627Z digest=sha256:6074f3e29773f5dbbee6176b21b46b678d4a6e7f32d057a568c0fa7a088ec80f

Observation bfcec33e-e2d4-4177-b44e-ca4ba28466f4 · outbound

This paper cites Generalists vs. Specialists: Evaluating LLMs on Highly-Constrained Biophysical Sequence Optimization Tasks.

Large Language Model is Secretly a Protein Sequence Optimizer Generalists vs. Specialists: Evaluating LLMs on Highly-Constrained Biophysical Sequence Optimization Tasks

Reference 4

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unresolved
no resolver link, observed 2026-08-10T20:10:52.809550Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.809550Z digest=sha256:5745234c6708177f35ec59367c1dbf1d5a3ba98306adb371995835a6037108c1

Observation bfce81a8-eb0f-48ac-b90b-6f9583c4c82c · outbound

This paper cites Flip: Benchmark tasks in fitness landscape inference for proteins.

Large Language Model is Secretly a Protein Sequence Optimizer Flip: Benchmark tasks in fitness landscape inference for proteins

Reference 5

Resolution
unresolved
no resolver link, observed 2026-08-10T20:10:52.813340Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.813340Z digest=sha256:c43802a5376a164c7f2c9b36e887fe866a9a326ab0278132482119ef8d0e8452

Observation da4c3c52-0da3-4c2f-b777-c74dba3c24cf · outbound

This paper cites Deep mutational scanning: a new style of protein science.

Large Language Model is Secretly a Protein Sequence Optimizer Deep mutational scanning: a new style of protein science

Reference 6

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.230322Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.816330Z digest=sha256:38acff3392775382f0cd6f93aaba83377d164490dbd18a046c878a2875b13435

Observation f3a784c4-729c-48af-adc8-51bab59e9985 · outbound

This paper cites Mutation effects predicted from sequence co-variation.

Large Language Model is Secretly a Protein Sequence Optimizer Mutation effects predicted from sequence co-variation

Reference 7

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.218767Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.819358Z digest=sha256:eb23e9d6ced59b00c72dd4e7ecfe2bae6fb86187cdb365efe16a0c35ed424ea8

Observation 03d5914f-99ce-4572-a099-c7dd7aa51904 · outbound

This paper cites Biological sequence design with gflownets.

Large Language Model is Secretly a Protein Sequence Optimizer Biological sequence design with gflownets

Reference 8

Resolution
unresolved
no resolver link, observed 2026-08-10T20:10:52.823344Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.823344Z digest=sha256:dab9ba1a581102c3e0851c19b62098467d95c337516aced79db130aac0aa2181

Observation e71a733a-283c-4c1c-9a10-2e5cd0f163a1 · outbound

This paper cites A combinatorially complete epistatic fitness landscape in an enzyme active site.

Large Language Model is Secretly a Protein Sequence Optimizer A combinatorially complete epistatic fitness landscape in an enzyme active site

Reference 9

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.199633Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.827108Z digest=sha256:f0fa651dd72f455f621d8d5eae8be8c49d2ce1f98e01e271bccfcf54b846874b

Observation 5e575411-3870-4459-ab6c-6c62e05e3555 · outbound

This paper cites Highly accurate protein structure prediction with alphafold.

Large Language Model is Secretly a Protein Sequence Optimizer Highly accurate protein structure prediction with alphafold

Reference 10

Resolution
unresolved
no resolver link, observed 2026-08-10T20:10:52.831447Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.831447Z digest=sha256:b7206ded1eba0ccf1e1702b2deeb27dbe9f5c32255e0f38505f862f0d0b90002

Observation bebdca3e-7445-4ade-9fc0-2c230a62830e · outbound

This paper cites Improving protein optimization with smoothed fitness landscapes.

Large Language Model is Secretly a Protein Sequence Optimizer Improving protein optimization with smoothed fitness landscapes

Reference 11

Resolution
unresolved
no resolver link, observed 2026-08-10T20:10:52.835278Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.835278Z digest=sha256:be18fb067a5165a8dfee485ae8156f2b84b33fc46b7be474c72632386b4b81a2

Observation 2f33d543-121c-458c-a8ec-5b6f2f588d8e · outbound

This paper cites Evolutionary-scale prediction of atomic-level protein structure with a language model.

Large Language Model is Secretly a Protein Sequence Optimizer Evolutionary-scale prediction of atomic-level protein structure with a language model

Reference 12

Resolution
unresolved
no resolver link, observed 2026-08-10T20:10:52.839155Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.839155Z digest=sha256:2d0f504b9c1b3a3567316ec798bf830e88e367bb11bf42fad8095c3f9048f821

Observation d7f02d13-fbda-4b6b-9e51-345bfc29aea7 · outbound

This paper cites Generative Design of Functional Metal Complexes Utilizing the Internal Knowledge of Large Language Models.

Large Language Model is Secretly a Protein Sequence Optimizer Generative Design of Functional Metal Complexes Utilizing the Internal Knowledge of Large Language Models

Reference 13

Resolution
verified exact
local_arxiv, observed 2026-08-10T20:10:53.040056Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.843234Z digest=sha256:1dbe2b05d0e6fdc88ab3e8c5f95420e348eec383aad3b2ce096622a2fc89a210

Observation 2f3782b1-65ac-4a13-b571-bc327ead024a · outbound

This paper cites Proximal exploration for model-guided protein sequence design.

Large Language Model is Secretly a Protein Sequence Optimizer Proximal exploration for model-guided protein sequence design

Reference 14

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.169929Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.847535Z digest=sha256:2dd4095c0725c329b63945cbe3e179a0ee3140b03ea696a2e1dea9f2873a18fb

Observation 2572dddf-6404-4a48-9222-841e884fb64b · outbound

This paper cites Exploring protein fitness landscapes by directed evolution.

Large Language Model is Secretly a Protein Sequence Optimizer Exploring protein fitness landscapes by directed evolution

Reference 15

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.159588Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.850788Z digest=sha256:e9675988895b07c831a106933c584235262e109ef2885ed663ce68a105f0cc86

Observation 979edd04-8f67-4ac1-9765-ade5f6571282 · outbound

This paper cites Local fitness landscape of the green fluorescent protein.

Large Language Model is Secretly a Protein Sequence Optimizer Local fitness landscape of the green fluorescent protein

Reference 16

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.150471Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.854264Z digest=sha256:eb6cc4246a071d29cd75f94bc51d5e89d3714e812265b570330ab3dd09396250

Observation 6712293c-a35f-45d0-9090-cf622aa36151 · outbound

This paper cites Tuned fitness landscapes for benchmarking model-guided protein design.

Large Language Model is Secretly a Protein Sequence Optimizer Tuned fitness landscapes for benchmarking model-guided protein design

Reference 17

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.139117Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.858978Z digest=sha256:5b3cca49ae7d8e73b9dc554f1802e0145d6febc871b2cbc1c0a03cee378859d2

Observation 45d721af-c36f-4066-acb0-4c7c3459f28d · outbound

This paper cites Protein design by directed evolution guided by large language models.

Large Language Model is Secretly a Protein Sequence Optimizer Protein design by directed evolution guided by large language models

Reference 18

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.127024Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.862125Z digest=sha256:9fde159620dcc4cce7247fb11864c92534e61c24b57db768795a51b34b8161a9

Observation 7de8ead5-dc67-4776-a252-91815a504cae · outbound

This paper cites Efficient Evolutionary Search Over Chemical Space with Large Language Models.

Large Language Model is Secretly a Protein Sequence Optimizer Efficient Evolutionary Search Over Chemical Space with Large Language Models

Reference 19

Resolution
unresolved
no resolver link, observed 2026-08-10T20:10:52.865174Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.865174Z digest=sha256:233c36ff74df8724af4aa60193a93c55283e293bf6aabc596000c2e77f46b51c

Observation 82fba977-7756-461a-9432-e3bfbf72b613 · outbound

This paper cites Adaptation in protein fitness landscapes is facilitated by indirect paths.

Large Language Model is Secretly a Protein Sequence Optimizer Adaptation in protein fitness landscapes is facilitated by indirect paths

Reference 20

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.116023Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.868848Z digest=sha256:babb3a74ccaf81f644a25bd04a088229b29664f58606c06f191efd5132ea9e34

Observation 15c1e4a0-a150-4a78-b959-fc264ef04db8 · outbound

This paper cites Active learning-assisted directed evolution.

Large Language Model is Secretly a Protein Sequence Optimizer Active learning-assisted directed evolution

Reference 21

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.103645Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.872396Z digest=sha256:0515ce46b0a7be680c6eb968b090f1ace21ef3ec8f1e281f58763610d1676abb

Observation 73277fba-128a-4273-8c10-98df0a423ddd · outbound

This paper cites Machine-learning-guided directed evolution for protein engineering.

Large Language Model is Secretly a Protein Sequence Optimizer Machine-learning-guided directed evolution for protein engineering

Reference 22

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.091723Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.875673Z digest=sha256:c02ad60b07bbdb084ba0e5d13fc3e77f3e44fcb2f6f402d050d71cf3e9ff40b0

Observation 612f45ef-1736-420f-952d-b57f949b0793 · outbound

This paper cites Protein language models learn evolutionary statistics of interacting sequence motifs.

Large Language Model is Secretly a Protein Sequence Optimizer Protein language models learn evolutionary statistics of interacting sequence motifs

Reference 23

Resolution
verified fuzzy
raw_fallback, observed 2026-08-10T20:10:53.079923Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-08-10T20:10:52.878940Z digest=sha256:7cb42bb71afa3a67abbf6fc710c20ba79948895e146921cdba277610d7f0d769

Observation 8266c117-45f1-4cd9-b727-1840f779eaaa · outbound

This paper cites write newline.

Large Language Model is Secretly a Protein Sequence Optimizer write newline

Reference 24

Resolution
unresolved
no resolver link, observed 2026-08-10T20:10:52.882094Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.882094Z digest=sha256:e6cb33bfc558e67c84b50207cdb93ff12e76565530355ecc7958b923cdf4ade1

Observation 4667a5e4-e4c8-4e74-a820-02d376f78519 · outbound

This paper cites @esa (Ref.

Large Language Model is Secretly a Protein Sequence Optimizer @esa (Ref

Reference 25

Resolution
unresolved
no resolver link, observed 2026-08-10T20:10:52.886231Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.886231Z digest=sha256:a04780ccb8d4884177ae8ff3476ea2dbeeced16ebe26e4b892528c0149f541d8

Observation 79807b36-d894-4383-a751-c3b3ec1ce17c · outbound

This paper cites an unresolved cited work.

Large Language Model is Secretly a Protein Sequence Optimizer Unresolved cited work

Reference 26

Resolution
unresolved
no resolver link, observed 2026-08-10T20:10:52.889790Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.889790Z digest=sha256:d05702292dd723c77fd0e8ed8b63fcc293dbc93c8458d7bd0c107c75fe9746db

Observation 51dce926-717b-48f5-a6c0-2f992ca2ea2c · outbound

This paper cites an unresolved cited work.

Large Language Model is Secretly a Protein Sequence Optimizer Unresolved cited work

Reference 27

Resolution
unresolved
no resolver link, observed 2026-08-10T20:10:52.893429Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=arxiv_source observed=2026-08-10T20:10:52.893429Z digest=sha256:59bed75b35113304e1f3f4bed3c9f91cb77652b7a7ddb077503a3288e0e8f7f8

Pith citing papers

Observation 711d76bf-83f7-403e-8a8f-1bf839ec8eb9 · inbound

Towards an AI co-scientist cites this paper.

Towards an AI co-scientist Large Language Model is Secretly a Protein Sequence Optimizer

Reference 279

Resolution
verified exact
arxiv_id, observed 2026-05-11T13:02:45.541768Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-05-11T13:02:43.571234Z digest=sha256:b26f7c0f1085d900f71956bd9e854645f24ce1290f2c10254d1c73b97cf2778d

Observation e2c469a8-6acf-486b-8f5e-d317bad7e637 · inbound

A Systematic Survey on Large Language Models for Evolutionary Optimization: From Modeling to Solving cites this paper.

A Systematic Survey on Large Language Models for Evolutionary Optimization: From Modeling to Solving Large Language Model is Secretly a Protein Sequence Optimizer

Reference 216

Resolution
unresolved
no resolver link, observed 2026-08-04T20:55:41.916450Z

Source-reported events for the cited work

Unavailable: canonical work link unavailable.

source=pdf_text observed=2026-08-04T20:55:41.916450Z digest=sha256:43f80c8b7264c1678ba5160b64ce5fadf4bdc95d608778bcc796acc15306edfd

Observation a062f0c5-ff33-46bf-8fcf-bfd9ee9b6403 · inbound

Towards Diverse Scientific Hypothesis Search with Large Language Models cites this paper.

Towards Diverse Scientific Hypothesis Search with Large Language Models Large Language Model is Secretly a Protein Sequence Optimizer

Reference 47

Resolution
verified exact
arxiv_id, observed 2026-07-03T04:07:36.966106Z

Source-reported events for the cited work

No event found in the named queried sources as of 2026-08-20T06:33:59.587034+00:00.

source=arxiv_source observed=2026-06-27T14:09:31.837231Z digest=sha256:a47a70574b988a9a38f1484479b932e76172c28dbbcf79cb67cc8d8ac8ca1805