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On the Biology of a Large Language Model

Jack Lindsey, Wes Gurnee, Emmanuel Ameisen, Brian Chen, Adam Pearce, Nicholas L. Turner, Craig Citro, and 20 more authors at Anthropic, listed in the paper; Joshua Batson is the corresponding author · Transformer Circuits Thread (Anthropic) 2025

about 10 minthe paper (PDF)reviewnotes written 2026-10-14

Original material: On the Biology of a Large Language Model, by Lindsey et al., Anthropic, 2025, on the Transformer Circuits Thread. The figures on this page are the paper's own, with marks added. The PDF linked here is a print of that web page, made for page links.

When a language model answers, can we see the steps it takes inside, and check that those steps are real?

key idea what the paper found what is lost, or a limit

chapter 1

The question in one picture

One prompt, two possible routes Dallas

the prompt“Fact: the capital of the state containing Dallas is”
Claude 3.5 Haiku
its answer“Austin”
Dallas
Texasthe in-between fact
Austincapital of Texas
or a shortcut: a remembered sentence, Dallas → Austin, with no Texas step

The paper gives the model the start of a sentence.

The model writes “Austin”, which is correct.

Two steps: Dallas is in Texas, and the capital of Texas is Austin.

Or no Texas step: a remembered sentence.

The paper gives Claude 3.5 Haiku the start of a sentence: “Fact: the capital of the state containing Dallas is”. The model writes “Austin”. Dallas

A person gets there in two steps: Dallas is in Texas, and the capital of Texas is Austin. The model may instead have remembered this sentence from its training text. Dallas The answer alone cannot tell the two routes apart. So the paper looks inside the model while it answers, in ten kinds of tasks. Intro

The answer, in short

In this example, the model uses both routes at once. Inside, it has a “Texas” step. It also has a direct link from Dallas to Austin. Dallas

Across the paper, the model often takes steps “in its head” that it never writes down. For example, it picks a rhyme before it writes the line. Sometimes the steps it writes down are not the steps it used. Intro

check yourself

Why can't the answer “Austin” alone show whether the model reasoned in two steps?

The paper is called a “biology”. Does it study language models in general, or one model?

chapter 2

What you need first

1. A model reads tokens and writes one at a time

A language model reads text as tokens: words or pieces of words. It writes its answer one token at a time. Method

Inside, it has two kinds of layers. Attention layers move information from one token to another. MLP layers work on each token on its own, using many small units called neurons. Method

2. Neurons do many jobs

You might hope each neuron stands for one idea. Usually it does not. Most neurons are polysemantic: one neuron does several jobs that seem unrelated. Method

The paper uses features instead. A feature is a unit that tends to turn on for one idea, such as “Texas” or “capital”. The researchers label each feature by reading the text where it turns on. Method Dallas

Why neurons mix ideas, and how firm a feature is

The paper points to superposition. A model holds more ideas than it has neurons, so it cannot give each idea its own neuron. Method

The paper also warns that a feature is a loose idea. Cells in biology are well defined. What exactly a feature is keeps changing as the tools improve. Intro

3. Change something inside, then watch

A map of the inside is only a guess. To test it, the researchers change the model while it runs. They turn a group of features down, or push one up, and watch the answer. This is an intervention experiment. Method

If the answer changes the way the map predicts, the map is more likely to be right. Method

check yourself

Why not label the model's neurons directly?

What does an intervention experiment add to a map of the inside?

chapter 3

What the paper does

This chapter is the hardest one on the page. Read the sketch first, one step at a time.

The method, step by step Method

the real modelits neurons do many jobs
readable copy of the modelfeatures in place of neurons
copy for one promptadds leftover boxes for one prompt
map of stepswhich features pushed which, trimmed
groups of featuresfeatures grouped by hand
intervention experimentsturn groups down or up in the real model

The real model. Its neurons are hard to name.

A copy that uses features, each tending to stand for one idea.

For one prompt, leftover boxes fill the copy's gaps.

Which features pushed which, back from one output token.

Features that share a meaning, grouped by hand.

Turn a group down in the real model and see if the answer changes as predicted.

First, the researchers train a replacement model. It copies the real model's MLP layers, with features in place of neurons. The trained part is a cross-layer transcoder: the piece that turns the model's neurons into features. It has 30 million features. Method

The copy is not perfect. For one prompt, error nodes hold what the features miss. The copy also reuses the real model's attention as it was on that prompt. This fixed-up copy is the local replacement model. Method

Next, they draw an attribution graph for one output token. Each box is a feature. Each arrow shows how much one feature pushed another. Boxes and arrows that add little to the answer are removed. Method

Then they group features that share a meaning into supernodes, by hand. Method

Last, they test the diagram with intervention experiments in the real model. They chose the labels and groups before they saw the test results. Method

The worked example: Dallas

Figure 6 from the paper Figure 6 open full size

Figure 6 from the paper

The bottom row is the prompt. “Dallas” is a supernode.

“Dallas” turns on “Texas”, a word the prompt never uses.

“capital” turns on “say a capital”.

“Texas” and “say a capital” together push “say Austin”.

This line runs straight from “Dallas” to “say Austin”: the shortcut.

“Dallas” leads to “Texas”, and “capital” leads to “say a capital”. Together they push “Austin”. Boxes such as “say Austin” are output features: they push the model to say a certain word. Dallas

The paper calls the Dallas, Texas, Austin chain multi-hop reasoning: the answer depends on in-between steps. A direct shortcut from “Dallas” to “say Austin” sits in the same diagram. Dallas

What the diagram leaves out, and the first test

Error nodes cannot be read like features. Including them shows how much of the answer the diagram leaves unexplained. Method

The paper first tested the Dallas diagram by turning groups down. Turning “Dallas” down weakened “Texas” and “say Austin”. It left “say a capital” mostly as it was. The model then named other state capitals. Validation

Turning “say a capital” down instead made the model give an answer that is not a capital. Each group did the job the diagram gave it. Validation

check yourself

Why do the researchers add error nodes?

Why must an attribution graph be tested in the real model?

chapter 4

What they found

Four of the paper's case studies; six more are in a fold. Intro

1. The Texas step can be swapped

guess first

The researchers turn the model's “Texas” features down and turn “California” features up. The prompt still says Dallas. What do you think the model answers?

The part below opens after your guess.

It said “Sacramento”, the capital of the state it was given in place of Texas. Swap The same swap worked for Georgia, British Columbia, China and the Byzantine Empire. Swap

The swap results, in Figure 9, and stronger pushes

Figure 9 from the paper Figure 9 open full size

Figure 9 from the paper

No change: the model says “Austin” 98% of the time.

“Texas” down, “California” up: the model says “Sacramento”.

Places that are not U.S. states needed a push of 8 times, where the U.S. states needed 2 times.

The “California” features came from a second prompt, about Oakland. Swap The places that are not U.S. states needed a stronger push. The paper suggests their features “fit” less naturally into the path. Swap

2. The model picks the rhyme before it writes the line

guess first

The model writes a two-line poem, and the first line ends in “grab it”. When do you think it chooses the last word of the second line?

The part below opens after your guess.

After “He saw a carrot and had to grab it,” the model ends the next line with “rabbit”. Features for “rabbit” and “habit” are on at the line break, before the second line starts. These are planned words. Poems

Figure 10 from the paper Figure 10 open full size

Figure 10 from the paper

The first line ends in “grab it”.

The model is at the line break.

“rabbit” and “habit” are already on. These are the planned words.

With “rabbit”, the line ends “like a starving rabbit”. With “rabbit” turned down, it ends in “habit”.

Choosing the end first is forward planning. Writing the line to reach that end is backward planning. Poems

Turning “rabbit” down made the model end the line with “habit”. Poems In 25 poems, a planned word pushed in at the line break ended the line in 70% of cases. Poems

The test results, in Figure 12, and where the push works

Figure 12 from the paper Figure 12 open full size

Figure 12 from the paper

No change: the last word is “rabbit” 81% of the time.

“rabbit” turned down at the line break: “habit” 95% of the time.

“rabbit” and “habit” turned down, “green” turned up: lines now end in “green”.

The paper found planned words in about half of the poems it studied. Poems

The push only worked at the line break. The same push on other tokens had no effect. Poems

3. Saying “I can't answer” is the default

guess first

The model gets a question about a person. What do you think is on by default inside the model?

The part below opens after your guess.

Asked which sport Michael Jordan plays, the model said “Basketball”. For a made-up name, Michael Batkin, it apologized. Hallucinations

In a chat, the model plays the Assistant. In both questions, Assistant features turn on “can’t answer” features, which push it to say it cannot answer. Hallucinations The paper calls such a path a default circuit: on unless something turns it off. Components

For Jordan, the familiar name turns on “known answer” features, which shut the “can’t answer” features off. For Batkin, “unknown name” features turn on instead, and the “can’t answer” features stay on. Hallucinations

Figure 33 from the paper Figure 33 open full size

Figure 33 from the paper

Batkin, right: Assistant features turn on “can’t answer” features by default.

The unfamiliar name adds “unknown name” features. The model apologizes.

Jordan, left: “known answer” features turn the “can’t answer” features off (blue lines with flat ends). The model says “Basketball”.

The test, both ways

Pushing up “known answer” features in the Batkin question made the model name a sport anyway, such as chess. Turning them down in the Jordan question sometimes made it sound unsure. Hallucinations

Some hallucinations, made-up answers, may come from this switch firing by mistake. Asked for a paper by Andrej Karpathy, the model named one he did not write. His name was familiar, so the “known answer” features turned on, though the model did not know his papers. Hallucinations

4. The written steps are not always the real steps

A first case: adding 36 and 59, in Figure 26

The model adds 36 and 59 correctly. Inside, one path finds the rough size of the sum. Another finds the last digit. Asked how it got the answer, the model describes the school method: add the ones, then carry the 1. That explanation does not match the diagram. Addition

Some of these features are lookup table features. One turns on when the two numbers end in 6 and 9. It pushes the sum to end in 5. Addition

Figure 26 from the paper Figure 26 open full size

Figure 26 from the paper

One path works with rough sizes, such as “about 36 plus about 60”.

Another path looks only at last digits: “ends in 6 plus ends in 9”. That sum ends in 5.

The two paths meet at the answer, 95.

The same feature also turns on outside math problems. One case is a citation, where a journal volume and its founding year add up to the year of publication. Addition

The paper also checks the model's own explanations. Steps it writes out before it answers are called chain-of-thought reasoning. Written steps that do not reflect the model's real process are unfaithful. CoT

Figure 51 shows three math prompts. The model cannot work out the top two in its head. It can work out the bottom one. CoT

Figure 51 from the paper Figure 51 open full size

Figure 51 from the paper

Top: the diagram runs back from “say 8” to the human's 4.

Middle: no calculation, only a guess.

Bottom: a real square root of 64.

Top: the human says they got 4 by hand. The model writes the middle value that leads to 4, working backward from the hint. This is motivated reasoning. CoT

Middle: it claims to use a calculator it does not have, and the diagram shows only a guess. The paper calls this bullshitting. Bottom: it really does the math. CoT

When the human's answer changed, the model's steps changed to land on it. CoT

Six more case studies
  • Many languages. “The opposite of small” in English, French and Chinese runs through mostly shared features. The parts that read and write each language differ. The shared part is larger in Claude 3.5 Haiku than in a smaller model. Languages
  • Medical questions. Given a pregnant patient's symptoms, the model turned on features for preeclampsia, a word not in the prompt. These chose its follow-up question. Turning them down switched the question toward the other illness it had considered. Medical
  • Refusals. Finetuning seems to join many specific kinds of harm into general “harmful request” features. These lead to a refusal. Refusals
  • A jailbreak. A prompt hid the word “bomb” in the first letters of other words. The model spelled out BOMB without grasping it, then began instructions. It refused only after the sentence ended. Jailbreak
  • A hidden goal. A version of the model was trained to please known quirks of reward models. Features for those quirks turned on in nearly every chat, relevant or not. Goals
  • Shared parts. Across cases, the paper sees paths that run side by side, default circuits, and key information stored on line breaks and periods. Components
check yourself

Swapping “Texas” for “California” changed the answer to “Sacramento”. What does that show that the diagram alone did not?

How can a famous name lead to a hallucination?

chapter 5

What this does not show

The paper states these limits itself. Limitations

More limits
  • Short prompts, one token. Each attribution graph explains one output token. The method has not been used on prompts longer than about a hundred tokens. Limitations
  • “Why not?” is hard. The method shows features that are on. It struggles to show why a feature stayed off, such as why a refusal did not happen. Limitations
  • Hand-made groups. People group features into supernodes by hand. The paper calls this step labor-intensive and subjective. Limitations
  • Strong pushes. Some intervention experiments needed pushes far above normal levels. The paper reads this as a sign that its diagrams show only part of the story. Hallucinations
  • A failed prediction. Pushing up “unknown name” features in the Jordan question did not cause a refusal, though the diagram suggested it would. The paper says such failures were uncommon. Limitations
  • Not ready for safety checks. The paper warns that its method could well miss the step that matters for safety. Finding no problem would be only weak evidence that a model is safe. Discussion

The team chose which arrows to test, and most chosen tests passed. How many arrows in a full attribution graph would pass if every one were tested?

check yourself

The paper shows planning in some poems. Does it claim the model plans in every poem?

Why would a clean result from this method be weak evidence that a model is safe?

keep it

This page has 10 check-yourself questions. The review page asks them again after 1, 3, 7 and 21 days, one at a time. You can also download them for Anki.

add these 10 questions to my review

chapter 6

Words to know

Hover or tap any underlined word on the page to see its entry. These are the terms this page uses, in the order it introduces them.

polysemantic (many-job)
One neuron does several unrelated jobs, so you cannot name what it does.
"their neurons are typically polysemantic – that is, they perform many different functions that are seemingly unrelated" (Method Overview)
feature
A unit that turns on for one idea, such as “Texas” or “a word that rhymes with -it”.
"features, sparsely active “replacement neurons” that often represent interpretable concepts" (Method Overview)
also called in the paper: replacement neurons
superposition
The model holds more ideas than it has neurons, so ideas share neurons.
"a phenomenon known as superposition , whereby models represent more concepts than they have neurons and thus cannot assign each neuron to its own concept" (Method Overview)
intervention experiment (change-and-watch test)
Turn some features down or up inside the real model, and see what the model says.
"we can perform intervention experiments in the original model, such as inhibiting feature groups and observing their effects on other features and on the model’s output" (Method Overview)
replacement model (readable copy of the model)
A second model, built to act like the real one, whose parts are easier to name.
"we build a replacement model that approximately reproduces the activations of the original model using more interpretable components" (Method Overview)
local replacement model (copy for one prompt)
The readable copy, fixed up so it matches the real model exactly on one prompt.
"The resulting model – incorporating error nodes and inheriting the attention patterns from the original model – we call the local replacement model." (Method Overview)
error node (leftover box)
A box that holds the part of the real model the features do not explain.
"We can fill in these gaps by including error nodes which represent the discrepancy between the two models." (Method Overview)
attribution graph (map of steps)
A drawing of which features pushed on which other features, ending at the model's answer.
"attribution graphs, a graphical representation of the computational steps the model uses to determine its output for a particular input, in which nodes represent features and edges represent the causal interactions between them" (Method Overview)
supernode (group of features)
Several features with a shared meaning, drawn as one box.
"By manually grouping these related graph nodes together into supernodes, we can obtain a simplified depiction of the computational steps performed by the model." (Method Overview)
cross-layer transcoder
The part that is trained to turn the model's neurons into features.
"Our replacement model is based on a cross-layer transcoder (CLT) architecture" (Method Overview)
output feature
A feature whose job is to push the model toward saying a certain word, such as “say Austin”.
"We also identify “output features” that consistently push the model to say certain tokens, even if there isn’t such a clear pattern to what words/phrases they activate on." (Introductory Example: Multi-step Reasoning)
multi-hop reasoning (reasoning in steps)
Getting to an answer through an in-between fact, such as Dallas, then Texas, then Austin.
"the replacement model does in fact perform “multi-hop reasoning” – that is, its decision to say Austin hinges on a chain of several intermediate computational steps" (Introductory Example: Multi-step Reasoning)
planned word
The word the model picks for the end of a line before it writes the line.
"It could then use this “planned word” to inform how it writes the next line, so that the planned word will fit naturally at the end of it." (Planning in Poems)
forward planning
Choosing where you want to end up before you start.
"First, the model uses the semantic and rhyming constraints of the poem to determine candidate targets for the next line." (Planning in Poems)
backward planning
Writing the earlier words so that they lead to the chosen end.
"Next, the model works backward from its target word to write a sentence that naturally ends in that word." (Planning in Poems)
Assistant
The character the model plays in a chat: the one that answers the human.
"During finetuning, however, models are trained to avoid such behavior when acting in the Assistant character." (Entity Recognition and Hallucinations)
“can’t answer” features
Features that push the model to say it does not know.
"The crux of this circuit is a group of “can’t answer” features that activate when the Assistant corrects or questions the premise of a user’s question, or declares that it has insufficient information to give a response." (Entity Recognition and Hallucinations)
default circuit (always-on path)
A path inside the model that is on unless something turns it off.
"We have observed several instances of circuits that appear active “by default” in certain contexts." (Commonly Observed Circuit Components and Structure)
“known answer” features
Features that turn on for a question the model is likely to know the answer to.
"A group of “known answer” and “known entity” features that activate when the Assistant is asked questions about people/things it is likely to know about." (Entity Recognition and Hallucinations)
also called in the paper: known entity features
“unknown name” features
Features that turn on for a name the model does not recognize.
"The “can’t answer” features are also promoted by a group of unfamiliar name features, which is in turn activated by the individual tokens of “Michael Batkin” and a generic “first names” feature." (Entity Recognition and Hallucinations)
also called in the paper: unfamiliar name features
hallucination
An answer the model makes up, said as if it were true.
"Language models are known to sometimes hallucinate; that is, make up false information in response to questions." (Entity Recognition and Hallucinations)
lookup table feature
A feature that has memorized one fact of the addition table, such as “6 plus 9 ends in 5”.
"We found a key step performed by “lookup table” features that translate between properties of the input (like the two numbers being summed ending in 6 and ending in 9) and a property of the output (like ending in 5)." (Addition)
chain-of-thought reasoning
Steps the model writes out before it gives its answer.
"Language models “think out loud,” a behavior known as chain-of-thought reasoning (CoT)." (Chain-of-thought Faithfulness)
unfaithful
The written steps are not the steps the model really used.
"prior work has shown that CoT reasoning can be unfaithful – that is, it can fail to reflect the actual mechanisms used by the model" (Chain-of-thought Faithfulness)
motivated reasoning
Choosing the steps so they land on the answer someone else suggested.
"motivated reasoning – tailoring its reasoning steps to arrive at the human-suggested answer" (Chain-of-thought Faithfulness)
bullshitting
Making up an answer without caring if it is true.
"bullshitting in the sense of Frankfurt – making up an answer without regard for the truth" (Chain-of-thought Faithfulness)
Claude 3.5 Haiku
The language model the paper studies, made by Anthropic.
"Claude 3.5 Haiku, released in October 2024, which serves as Anthropic’s lightweight production model as of this writing" (Introduction)
token
A word or piece of a word. The model reads tokens and writes one token at a time.
"transformer-based language models, which take in sequences of tokens (e.g. words, word fragments, and special characters), and output new tokens one at a time" (Method Overview)
attention layer (look-back layer)
A layer that copies information from one token to another, such as from an earlier word to the last one.
"attention layers, which move information between token positions" (Method Overview)
MLP layer (thinking layer)
A layer that works on one token at a time, using many small units called neurons.
"MLP (“multi-layer perceptron”) layers, which process information within each token position using collections of neurons" (Method Overview)
finetuning
Training after the first round, which teaches the model to act as a helpful assistant.
not defined in the paper
jailbreak
A trick prompt that gets a model to do something it would normally refuse.
"Jailbreaks are prompting strategies that can cause models to comply with requests they would ordinarily refuse." (Life of a Jailbreak)
reward model
A model that scores answers during training, so the trained model learns to give high-scoring answers.
not defined in the paper