MLP neurons in transformer language models are widely understood to be polysemantic, motivating transcoders - sparse, learned approximations of MLP layers - as interpretable replacement models for mechanistic analysis. However, recent work shows that MLP neurons can be used to recover sparse, interpretable circuits (Arora et al. 2026, Ye et al. 2025), raising the question of whether transcoders are necessary for circuit analysis. We define neuron features as transcoder features whose weights are strongly aligned with individual MLP neurons, and show that they account for a substantial fraction of active features in late layers across 13 open-source transcoders. Only neuron features have a stable correspondence with something in the underlying MLP: other features vary more between training runs, do not match the MLP's Jacobian, and change less significantly from their random initialization. Our findings reframe MLP neurons as a stronger basis for interpretability than commonly assumed, and call for better methods of attributing the non-neuron structure that is used by the model.
Figures & tables
Figure 2: A summary of findings from our evaluations and case studies.
Figure 3: Reconstruction and replacement (a) and sort eval monosemanticity (b) for neurons vs transcoders. Top neuron activations explain most of the model’s computation and are similarly interpretable to transcoder feature activations.
Table 3Figure 4Figure 5
Appendix figures & tables20 assets
Supplementary material from the paper’s appendix.
Appendix
Contents
Appendix A
Appendix A Model and transcoder details
Appendix B
Appendix B Evaluation methodology
Appendix C
Appendix C Full evaluation results
Appendix D
Appendix D Features not aligned to single neurons
Appendix E
Appendix E Neuron feature prevalence and coverage
Appendix F
Appendix F Panels of neuron and neuron feature co-activation
Appendix
Table 6
Model
HF slug
Layers
dmodel
dmlp
Activation
Dense
Gemma 2 2B
google/gemma-2-2b
26
2304
9,216
GeGLU
Gemma 3 1B
google/gemma-3-1b-pt
26
1152
6,912
GeGLU
Gemma 3 4B
google/gemma-3-4b-pt
34
2560
10,240
GeGLU
Gemma 3 12B
google/gemma-3-12b-pt
48
3840
15,360
GeGLU
Gemma 3 27B
google/gemma-3-27b-pt
62
5376
21,504
GeGLU
Appendix
Table 3: Model architectures used in this work. All models use gated MLPs ( up , gate , down ). GeGLU uses GELUtanh [ Hendrycks and Gimpel, 2016 ] for the gate, SiGLU uses SiLU. Llama and Qwen use standard pre-norm around each sublayer. Gemma 3 is unusual in additionally applying a post_feedforward_layernorm after the MLP; this requires dividing by the LayerNorm weight to recover a faithful encoder/decoder geometry ( Section D.1 ) and shows up, among other things, as higher KL on Gemma 3 transcoders compared to the matched Llama/Qwen ones at similar L0 (Figure 12 ). For the two mixture-of-experts models, dmlp is written as the shared expert’s width plus the number of routed experts times their width. Both models have 8 active experts per token. Gemma 4 26B-A4B is used for bigram probing ( Table 7 ) and for testing expert concentration in attribution graphs ( Appendix N ). Qwen 3.5 35B-A3B is the model we train mixture of experts transcoders on ( Appendix O ). Model families used: Gemma 2 [ Gemma Team, 2024 ] , Gemma 3 [ Gemma Team, 2025 ] , Gemma 4 [ Gemma Team, 2026 ] , Qwen 3 [ Qwen Team, 2025 ] , Qwen 3.5 [ Qwen Team, 2026a ] , Llama 3 [ Llama Team, 2024 ] .
Transcoder
Model
Act.
Width
L0
Alive@66%
NF@66%
NF% acts@66%
GS-small
Gemma 3 1B
JumpReLU
262k
45
193k ( 74% )
15k ( 5.9% )
57.9%
GS-big
Gemma 3 1B
JumpReLU
262k
148
212k ( 81% )
24k ( 9.1% )
57.5%
GS-small
Gemma 3 4B
JumpReLU
262k
47
175k ( 67% )
23k ( 8.9% )
63.8%
GS-big
Gemma 3 4B
JumpReLU
262k
154
200k ( 76% )
33k ( 12.7% )
66.6%
GS-small
Gemma 3 12B
JumpReLU
262k
47
165k ( 63% )
41k ( 15.6% )
73.7%
GS-small
Gemma 3 27B
JumpReLU
262k
41
194k ( 74% )
97k ( 37.1% )
82.8%
Appendix
Table 4: Models used in the analysis of open source transcoders. GS is Gemmascope on Gemma 3 ( google/gemma-scope-2-{1,4,12,27}b-pt ); mwhanna is mwhanna/qwen3-{1.7,4,8,14}b-transcoders ; LXTC-8x and LXTC-32x are OpenMOSS-Team/Llama3_1-8B-Base-LXTC-{8,32}x ; mntss is mntss/clt-gemma-2-2b-426k and mntss/clt-llama-3.2-1b-524k (cross-layer transcoders; the width in the table is the per-layer width and the width in the name is the maximum width of the cross-layer decoder). All metrics measured on FineWeb at layer ⌊2N/3⌋ (where N is the number of layers; at 66% of the way into the model). L0 = mean active features per token. Alive = features with fire_count>0 ; NF = features with encoder and decoder cosine similarity both above 0.6 to the same neuron; NF% acts = share of feature firings that come from neuron features. The last two rows are the skip transcoders mntss/skip-transcoders-gemma-2-2b and mntss/transcoder-Llama-3.2-1B , used for sparse probing ( Table 6 ). Neither release states its activation function; we measure TopK for the Llama one (FVU 0.15 against 0.99 for ReLU) and assume it for the Gemma one, which we do not measure further.
Figure 13: Gated-MLP neuron geometry. Per-neuron activations are heavy-tailed (left); high-magnitude activations split into two regimes that share the same gate-positive half-plane (centre); and within each regime the effective encoder direction concentrates along Wup(i)±Wgate(i) across neurons (right).
Model
Neurons
Transcoder
Δ
Llama 3.1 8B
0.856
0.845
+0.011±0.007
Gemma 3 4B
0.854
0.832
+0.022±0.008
Qwen 3 8B
0.851
0.806
+0.045±0.011
Qwen 3 4B
0.833
0.794
+0.039±0.011
Qwen 3 1.7B
0.811
0.762
+0.050±0.010
Gemma 3 1B
0.834
0.792
+0.042±0.011
Appendix
Table 5: Sparse probing mean AUC at k=1 across the 113 [ Kantamneni et al., 2025 ] tasks. Neurons : dense MLP neurons. Transcoder : best-available transcoder per model ( Table 4 ). Δ shows the per-task mean difference. Neurons match or exceed the transcoder across all models. Per-task and full k -sweep tables in Appendix C.1 .
Name
AUC, k=1
AUC, k=4
AUC, k=16
Tasks won ( k=1 )
Model
Dictionary
Neurons
Dictionary
Neurons
Dictionary
Neurons
Dictionary
Neurons
Dictionary
Ties
Llama 3.1 8B
LXTC-8 ×
0.883
0.843
0.908
0.888
0.925
0.911
79
15
19
Llama 3.1 8B
LXTC-32 ×
0.883
0.839
0.908
0.886
0.925
0.910
80
15
18
Gemma 3 4B
GS-big
0.877
0.849
0.899
0.888
0.919
0.907
70
18
25
Qwen 3 8B
mwhanna
0.876
0.829
0.902
0.877
0.921
0.900
83
13
17
Qwen 3 4B
mwhanna
0.864
0.819
0.891
0.866
0.909
0.890
79
15
19
Appendix
Table 6: Sparse probing results across nine (model, dictionary) pairs on the 113 -task suite of Kantamneni et al. [2025] . Neurons : post-GeGLU/SiLU sign-split neurons. Dictionary : canonical post-activation learned dictionary (Appendix B.1 ). Mean peak test AUC across the 113 tasks (peak = max over layers per task), then averaged across tasks; bold marks the higher of the two within each (k,row) . Tasks won columns: number of tasks where one method beats the other by >0.005 AUC at k=1 ; ties otherwise. Bold marks the larger of the two win counts per row. Win rates and AUC gaps narrow with k but the sign of the gap is preserved at every k on every model.
Gemma 3 4B
Qwen 3 4B
Gemma 4 26B-A4B
Neurons
Gemmascope
Neurons
Matryoshka TopK
Neurons
Neurons, routed
Layer
k=1
k=8
k=1
k=8
k=1
k=8
k=1
k=8
k=1
k=8
k=1
k=8
L0
.891
.991
.988
.996
.946
.995
.992
.999
.723
.895
.902
.983
L1
.886
.982
.977
.990
.955
.993
.987
.998
.844
.977
.957
.997
L2
.854
.987
.985
.995
.950
.992
.979
.997
.833
.983
.937
.986
L3
.875
.985
.984
.996
.949
.990
.977
.996
.783
.982
.929
.987
Appendix
Table 7: Per-layer mean test AUC on the 1,500 bigram probing tasks (Appendix B.2 ), at k∈{1,8} selected features per probe. Match reports the rate at which the chosen (best k=1 across L0–L5) feature’s top-1 max-activating example contains the target bigram. The transcoder wins every AUC comparison on every model, and its features match the bigram in their top-1 example far more often than sparsified neurons do. The MoE model, Gemma 4, is close to the some of the transcoders’ level of performance; this is preliminary evidence that scaled up MoE training could lead to similar monosemanticity to that seen in transcoders.
32 k
65 k
131 k
262 k
524 k
1 M
Our TopK pipeline
TopK k=64
32 k
65 k
130 k
259 k
—
895 k
TopK k=128
32 k
65 k
131 k
261 k
—
939 k
TopK k=32
32 k
64 k
126 k
247 k
—
754 k
JumpReLU pipeline
JumpReLU L0≈30
—
—
—
202 k
263 k
262 k
Appendix
Table 8: Effective dictionary size at width M , defined as the number of features with non-zero firing on a buffer of 100 steps ( ∼200 k tokens).
Figure 14
Dict.
Active
Model + transcoder
early
late
early
late
Gemma 3 1B L0-small
2.1
3.6
27
48 [ 1.8× ]
Gemma 3 1B L0-big
4.9
6.7
46
52 [ 1.1× ]
Gemma 3 4B L0-small
2.6
7.2
22
53 [ 2.4× ]
Gemma 3 4B L0-big
6.7
11.2
41
59 [ 1.5× ]
Gemma 3 12B small
2.5
10.7
12
43 [ 3.6× ]
Appendix
Table 9: Neuron features in 13 open-source transcoders, defined by encoder >0.7 and decoder >0.7 to the same neuron. Numbers are % of the dictionary ( Dict. ) and % of active features ( Active ), averaged separately over layers in the first vs. second half of each model. Brackets in the rightmost column give the ratio between the share in late and early layers. See Appendix A for transcoder details.
Threshold
32k
65k
128k
256k
1M
∣cos∣≥0.5
86.0
91.7
94.1
96.4
96.7
∣cos∣≥0.7
78.3
86.7
89.9
93.3
93.0
∣cos∣≥0.9
55.9
71.5
78.1
86.0
84.6
Appendix
Table 10: Percentage of Gemma 3 4B L22 MLP neurons matched by at least one TopK transcoder feature across dictionary size (columns) and cosine threshold (rows).
Layer
n
med. AUC
enc bin
n
med. Gini
5
35
0.911
[0.30,0.50)
29
0.224
10
37
0.555
[0.50,0.70)
15
0.319
15
38
0.873
[0.70,0.85)
35
0.956
20
31
0.998
[0.85,1.00]
119
0.996
25
28
0.993
30
36
0.983
Appendix
Table 11: Left: per-layer median AUC. Right: median Gini (rescaled AUC) by encoder cosine similarity: only features we call neuron features have high median discriminability.
L0=100
L0=25
L0=10
Model + transcoder
RS
early
mid
late
early
mid
late
early
mid
late
Gemma 3 1B small
0.62
23 ± 0
37 ± 1
46 ± 1
16 ± 0
33 ± 1
45 ± 1
13 ± 0
25 ± 1
51 ± 1
Gemma 3 4B small
0.56
25 ± 0
39 ± 0
60 ± 1
15 ± 0
29 ± 1
56 ± 1
13 ± 0
24 ± 1
60 ± 1
Gemma 3 12B small
0.50
19 ± 0
36 ± 1
62 ± 1
16 ± 0
37 ± 1
66 ± 1
13 ± 0
37 ± 1
75 ± 2
Gemma 3 27B small
0.49
22 ± 0
41 ± 1
70 ± 1
15 ± 0
30 ± 1
72 ± 2
11 ± 0
26 ± 2
77 ± 3
Gemma 2 2B mntss
0.66
17 ± 0
44 ± 1
49 ± 1
11 ± 0
34 ± 1
45 ± 1
10 ± 0
27 ± 1
43 ± 1
Appendix
Table 12: Percentage of neuron features in attribution graphs across nine open-source transcoders, by layer group and pruning strength. RS : median replacement score over circuits; lower for only the larger Qwen 3 transcoders. early / mid / late : percentage of neuron features in each of the three groups of layers.
Figure 43: “Red” case study on Gemma 3 4B L22. Top: attribution graph for “Translation of the word красное :” (predicting “Red”/“red”), with N5039 highlighted. Bottom: max-activating contexts for selected MLP neurons, with activations N7405 + =8.00 , N5039 + =5.31 , N10102 + =1.40 , N4137 + =0.9 , N10100 + =0.86 .
Figure 44: “Red” case study, k=64 TopK transcoder. Attribution graph shows two mildly (light orange; encoder alignment, from 0 to 1, is represented as color, from white to orange) neuron-aligned features and one strongly neuron-aligned feature (F138722). N10102, the neuron it is aligned to, activates on “red”/“ted” (§ 45(d) ). The other two features are weakly aligned to neuron 5039, and both activate only on the token “red”. Figure 46 shows that they both cluster the neuron.
Figure 45: “Red” case study, Gemmascope transcoder. Attribution graph shows three neuron-aligned features in dark orange. One feature is aligned to a color neuron, 7405. Another is aligned to N10102, a “red”/“ted” neuron, the activations of which are shown in the bottom right. The third one is a neuron feature for N5039 and shows the same polysemanticity between “red” and “transparent”.
Transcoder
p1
p2:4
Our transcoders
Baseline k=64
—
—
Baseline k=32
0.64
0.77
NI k=32
0.66
0.70
NFT k=32 , G=16,r=4
1.00
0.66
SNRL k=32
0.71
0.57
Appendix
Table 13: Absorption evaluation for different transcoder-based clustering methods on Gemma 3 4B L22, with top K=128 neurons considered active and 10 M SmolLM2 tokens. p1 is the probability the feature’s top-1 weight-aligned neuron is in the active set; p2:4 the same for the next three weight-aligned neurons.
Transcoder
FVU ↓
KL ↓
Jac. faith. ↑
Our transcoders
Baseline k=64
0.166
0.77
0.276
Baseline k=32
0.195
1.29
0.225
NI k=32
0.209
1.16
0.238
NFT k=32 , G=16,r=4
0.221
1.15
0.230
SNRL k=32
0.213
1.38
0.232
Appendix
Table 14: Reconstruction quality and Jacobian faithfulness on Gemma 3 4B layer 22. FVU and KL are over 1 M held-out SmolLM2 tokens. “Jac. faith.” is the cosine similarity between the MLP Jacobian and the transcoder Jacobian, divided by Layernorm for the Gemmascopes: the post-LN scaling factor (1+wln) is removed so the comparison lives in pre-LN MLP-output space. FVU, however, is measured against a different target for our transcoders and Gemmascopes.
Clustering
CM acc ↑
TNM acc ↑
Our transcoders
Baseline k=32
0.937±0.005
0.731±0.030
NI k=32
0.918±0.008
0.629±0.031
NFT k=32 , G=16,r=4
0.918±0.010
0.598±0.033
SNRL k=32
0.927±0.008
0.657±0.030
Gemmascope
Appendix
Table 15: Within-cluster sort eval results on 50 random Gemma 3 4B L22 neurons (judge: Claude Sonnet 4.6, 60 trials per neuron). Up to 10 neurons per transcoder produced too few viable clusters and were dropped (more for Gemmascope, which has sparser per-panel cluster coverage). CM acc = Cluster match accuracy (SE over panels), TNM acc = Top-neuron match accuracy (binomial SE over panel-cluster pairs).