ND4LNADH dehydrogenase subunit 4L

ND4L is the smallest Complex I subunit encoded by mtDNA, a membrane protein of about 100 amino acids. In vertebrates the end of its gene overlaps the start of ND4, and the two are translated from one bicistronic transcript.

Median length

297 bp

Range 234–408 bp

Median GC content

45.5%

Middle half 39.0%–48.8%

Species with data

2,335

Of 2,335 genomes

Strand

+ strand

In 2,283 of 2,335 genomes

ND4L in the mitochondrial genome of

Human Homo sapiens

RefSeq NC_012920.1 · 16,569 bp · GC 44.4% · ND4L at 10,470–10,766, 297 bp, + strand · Species page

ND4LProtein-coding gene

NADH dehydrogenase subunit 4L

Position: 10,470–10,766
Length: 297 bp
Strand: +
GC content: 43.1%

ND4L across all species

Protein-coding13
rRNA2
tRNA22
D-loop1

Outer ring: + strand, arrows run clockwise. Inner ring: − strand.
Middle ring: GC% of each gene against the whole genome (44.4%, dashed): out is GC-richer, in is AT-richer (full height = 6 points).

GC content of ND4L across classes

2,335 species, reference strand as deposited in RefSeq

Mammalsn 627 · median 37.7% · middle half 35.7%–40.1%

Birdsn 423 · median 47.8% · middle half 46.1%–49.2%

Ray-finned fishesn 895 · median 48.8% · middle half 46.8%–50.8%

Reptilesn 136 · median 40.0% · middle half 37.9%–42.4%

Sharks and raysn 87 · median 42.1% · middle half 40.1%–43.8%

Amphibiansn 47 · median 37.7% · middle half 34.4%–42.1%

Other classesn 120 · median 33.0% · middle half 28.6%–37.2%

Each dot is one species. Boxes span the middle half of the species (25th to 75th percentile) and the thick line marks the median; thin lines run from the lowest to the highest value.

Compare ND4L across taxa

GC content of ND4L against lifespan

One point per species, coloured by class

Hover or tap a point to see the species; select it again to open its page. Legend buttons spotlight a class.

Open in the plot

Correlation with maximum lifespan

All species, each class and each order with enough species

MeasureGC%LengthGC skewAT skew

GC content of ND4L against log₁₀ maximum lifespan in 32 groups; 8 pass q < 0.05. Plot it in the explorer

Correlation of GC content of ND4L with log10 maximum lifespan by group
Group	n	Pearson r	Spearman ρ	q	PIC r	Partial r
All speciesevery class	2,202	+0.00	−0.11	0.89	+0.00	+0.19
Ray-finned fishesclass Actinopterygii	827	+0.10	+0.05	0.012 (significant)	−0.07	−0.14
Mammalsclass Mammalia	615	+0.30	+0.31	< 0.001 (significant)	+0.07	+0.21
Birdsclass Aves	405	−0.04	−0.11	0.53	+0.03	+0.01
Reptilesclass Reptilia	126	+0.27	+0.29	0.022 (significant)	+0.06	+0.23
Sharks and raysclass Chondrichthyes	82	−0.02	−0.03	1.00	+0.12	−0.04
Amphibiansclass Amphibia	44	+0.15	+0.06	1.00	+0.17	–
Bivalvesclass Bivalvia	35	−0.28	−0.18	0.61	+0.16	–
Perciformesorder	208	−0.21	−0.34	0.008 (significant)	−0.04	−0.41
Passeriformesorder	140	+0.21	+0.20	0.057	+0.21	+0.14
Artiodactylaorder	134	+0.40	+0.27	< 0.001 (significant)	+0.06	+0.15
Cypriniformesorder	127	+0.29	+0.25	0.008 (significant)	+0.15	+0.19
Carnivoraorder	123	+0.27	+0.30	0.011 (significant)	+0.10	+0.27
Primatesorder	112	+0.36	+0.33	< 0.001 (significant)	+0.02	+0.07
Rodentiaorder	87	−0.16	−0.18	0.29	+0.08	−0.12
Squamataorder	60	+0.15	+0.24	0.77	+0.18	+0.30
Charadriiformesorder	51	+0.10	+0.06	0.76	+0.09	−0.07
Testudinesorder	49	−0.04	−0.11	0.95	−0.11	+0.05
Psittaciformesorder	44	−0.09	−0.12	0.76	+0.17	+0.07
Chiropteraorder	43	+0.22	+0.18	0.86	+0.24	+0.21
Clupeiformesorder	38	+0.01	+0.03	0.99	−0.30	–
Pleuronectiformesorder	35	+0.24	+0.12	0.39	−0.01	+0.19
Salmoniformesorder	35	+0.14	+0.10	0.77	−0.26	+0.16
Carcharhiniformesorder	31	+0.12	+0.24	0.74	+0.09	+0.07
Cetaceaorder	28	+0.24	+0.32	0.48	−0.00	−0.17
Anseriformesorder	25	−0.22	−0.22	0.60	−0.15	−0.03
Caudataorder	25	+0.32	+0.10	0.79	+0.08	–
Siluriformesorder	24	+0.27	+0.29	0.41	+0.18	–
Centrarchiformesorder	22	+0.43	+0.26	0.23	+0.33	–
Acipenseriformesorder	21	−0.32	−0.47	0.38	−0.31	−0.28
Rajiformesorder	21	−0.05	+0.14	0.96	+0.21	–
Scombriformesorder	21	−0.04	−0.15	0.98	−0.18	–

Length of ND4L against log₁₀ maximum lifespan in 11 groups; 2 pass q < 0.05. Plot it in the explorer

Correlation of Length of ND4L with log10 maximum lifespan by group
Group	n	Pearson r	Spearman ρ	q	PIC r	Partial r
All speciesevery class	2,202	+0.09	+0.01	< 0.001 (significant)	−0.01	+0.00
Ray-finned fishesclass Actinopterygii	827	+0.07	−0.01	0.061	+0.00	+0.02
Mammalsclass Mammalia	615	−0.01	+0.01	0.92	+0.02	+0.00
Reptilesclass Reptilia	126	+0.25	+0.17	0.031 (significant)	−0.06	+0.38
Amphibiansclass Amphibia	44	−0.06	−0.06	1.00	+0.03	–
Bivalvesclass Bivalvia	35	+0.12	+0.18	0.87	−0.22	–
Perciformesorder	208	−0.01	−0.02	0.94	−0.01	+0.09
Carnivoraorder	123	+0.03	+0.09	0.83	+0.04	+0.04
Primatesorder	112	−0.00	+0.01	1.00	+0.03	+0.06
Squamataorder	60	+0.03	−0.06	0.95	−0.13	+0.36
Testudinesorder	49	−0.43	−0.44	0.23	−0.12	−0.28

GC skew of ND4L against log₁₀ maximum lifespan in 32 groups; 7 pass q < 0.05. Plot it in the explorer

Correlation of GC skew of ND4L with log10 maximum lifespan by group
Group	n	Pearson r	Spearman ρ	q	PIC r	Partial r
All speciesevery class	2,202	−0.13	−0.15	< 0.001 (significant)	−0.01	−0.15
Ray-finned fishesclass Actinopterygii	827	−0.08	−0.12	0.032 (significant)	−0.04	+0.11
Mammalsclass Mammalia	615	−0.13	−0.13	0.003 (significant)	+0.02	−0.18
Birdsclass Aves	405	+0.29	+0.30	< 0.001 (significant)	+0.09	+0.14
Reptilesclass Reptilia	126	−0.20	−0.24	0.10	+0.01	−0.02
Sharks and raysclass Chondrichthyes	82	+0.01	−0.02	1.00	+0.48	+0.27
Amphibiansclass Amphibia	44	−0.10	−0.10	1.00	−0.06	–
Bivalvesclass Bivalvia	35	+0.10	+0.08	0.89	+0.04	–
Perciformesorder	208	+0.13	+0.18	0.14	−0.15	+0.58
Passeriformesorder	140	+0.16	+0.15	0.15	+0.24	+0.04
Artiodactylaorder	134	−0.12	+0.01	0.30	+0.21	+0.02
Cypriniformesorder	127	+0.12	+0.09	0.35	−0.12	−0.21
Carnivoraorder	123	+0.15	+0.15	0.19	−0.00	−0.15
Primatesorder	112	−0.30	−0.31	0.003 (significant)	+0.01	+0.01
Rodentiaorder	87	+0.28	+0.28	0.040 (significant)	+0.04	+0.14
Squamataorder	60	−0.08	−0.11	0.88	−0.15	−0.02
Charadriiformesorder	51	+0.19	+0.23	0.49	+0.04	+0.14
Testudinesorder	49	−0.14	−0.14	0.75	+0.10	−0.14
Psittaciformesorder	44	+0.08	+0.06	0.77	+0.21	−0.20
Chiropteraorder	43	−0.14	−0.18	0.90	−0.21	−0.13
Clupeiformesorder	38	−0.21	−0.28	0.66	+0.68	–
Pleuronectiformesorder	35	+0.08	+0.00	0.83	+0.17	−0.20
Salmoniformesorder	35	+0.09	+0.20	0.92	−0.00	+0.59
Carcharhiniformesorder	31	−0.05	−0.03	0.91	+0.01	+0.22
Cetaceaorder	28	−0.05	−0.11	0.89	−0.28	−0.11
Anseriformesorder	25	−0.05	−0.02	0.90	−0.13	+0.01
Caudataorder	25	−0.16	−0.27	0.83	+0.14	–
Siluriformesorder	24	−0.12	−0.15	0.75	−0.13	–
Centrarchiformesorder	22	−0.29	−0.30	0.42	−0.21	–
Acipenseriformesorder	21	−0.68	−0.50	0.013 (significant)	−0.71	−0.51
Rajiformesorder	21	−0.02	−0.00	0.98	+0.60	–
Scombriformesorder	21	−0.04	−0.02	0.98	−0.00	–

AT skew of ND4L against log₁₀ maximum lifespan in 32 groups; 6 pass q < 0.05. Plot it in the explorer

Correlation of AT skew of ND4L with log10 maximum lifespan by group
Group	n	Pearson r	Spearman ρ	q	PIC r	Partial r
All speciesevery class	2,202	+0.10	+0.16	< 0.001 (significant)	−0.13	+0.14
Ray-finned fishesclass Actinopterygii	827	+0.08	+0.05	0.045 (significant)	−0.18	−0.11
Mammalsclass Mammalia	615	+0.14	+0.14	< 0.001 (significant)	−0.00	−0.07
Birdsclass Aves	405	+0.04	+0.06	0.56	−0.08	+0.06
Reptilesclass Reptilia	126	−0.46	−0.47	< 0.001 (significant)	−0.09	−0.37
Sharks and raysclass Chondrichthyes	82	−0.06	−0.09	1.00	−0.60	−0.04
Amphibiansclass Amphibia	44	−0.06	−0.04	1.00	−0.03	–
Bivalvesclass Bivalvia	35	+0.03	+0.12	0.97	−0.04	–
Perciformesorder	208	+0.04	−0.00	0.70	+0.06	−0.31
Passeriformesorder	140	−0.23	−0.21	0.044 (significant)	−0.10	−0.01
Artiodactylaorder	134	−0.08	−0.14	0.46	−0.16	−0.16
Cypriniformesorder	127	+0.04	+0.07	0.75	+0.02	+0.07
Carnivoraorder	123	−0.00	+0.06	1.00	+0.02	−0.08
Primatesorder	112	+0.29	+0.30	0.004 (significant)	+0.04	+0.04
Rodentiaorder	87	−0.25	−0.27	0.069	+0.02	−0.09
Squamataorder	60	−0.26	−0.22	0.34	−0.07	−0.43
Charadriiformesorder	51	+0.08	+0.07	0.81	−0.11	−0.16
Testudinesorder	49	+0.13	+0.13	0.76	−0.11	+0.11
Psittaciformesorder	44	−0.14	−0.08	0.60	−0.08	+0.00
Chiropteraorder	43	−0.11	−0.06	0.90	+0.02	−0.18
Clupeiformesorder	38	+0.02	−0.05	0.98	−0.78	–
Pleuronectiformesorder	35	−0.41	−0.37	0.12	−0.30	+0.29
Salmoniformesorder	35	+0.16	+0.08	0.71	+0.17	−0.26
Carcharhiniformesorder	31	−0.26	−0.22	0.52	−0.04	−0.11
Cetaceaorder	28	+0.08	+0.08	0.82	+0.31	+0.27
Anseriformesorder	25	−0.12	−0.19	0.79	+0.16	+0.02
Caudataorder	25	−0.05	−0.02	0.89	−0.18	–
Siluriformesorder	24	+0.35	+0.37	0.26	+0.23	–
Centrarchiformesorder	22	+0.47	+0.40	0.23	+0.21	–
Acipenseriformesorder	21	−0.22	+0.23	0.63	−0.69	+0.30
Rajiformesorder	21	−0.46	−0.50	0.88	−0.71	–
Scombriformesorder	21	+0.13	+0.09	0.89	+0.01	–

n: species with both values in the group.
Pearson r, Spearman ρ: strength of the link with log₁₀ lifespan, from −1 to +1; Spearman uses ranks, so a few extreme species weigh less.
q: p-value adjusted for testing every mtDNA feature in the same group (Benjamini–Hochberg); rows with q < 0.05 are highlighted.
PIC r: the same test on phylogenetically independent contrasts, which removes similarity that comes from shared ancestry.
Partial r: the link with lifespan after accounting for body mass (log₁₀ adult weight), since bigger animals tend to live longer.

ND4L aligned across species

2,316 species, 193 columns; 98 columns have a residue in at least half of the species, and in 35 of those one residue is in 90% or more. MAFFT v7.526, FFT-NS-2 (progressive)

Loading the alignment of ND4L: 2,316 species, 193 columns.

All alignment files

Codon usage of ND4L

Vertebrates, vertebrate mitochondrial code

Most used codons

Share of all 218,438 sense codons of ND4L in 2,231 vertebrates

CUALeu9.2%
AUAMet5.2%
GCCAla4.8%
UUALeu4.0%
CUCLeu4.0%
GCAAla3.5%
UCASer3.2%
CUULeu3.2%

Preferred codon for each amino acid

Most used synonymous codon and its RSCU, pooled by class; darker cells show a stronger preference

Preferred codon and RSCU for each amino acid in ND4L, by class
Amino acid	All vertebratesn 2,231	Mammalsn 627	Birdsn 423	Ray-finned fishesn 895	Reptilesn 136	Sharks and raysn 87	Amphibiansn 47
PheF · 2 codons	UUC1.09	UUC1.06	UUC1.60	UUU1.03	UUC1.04	UUU1.09	UUU1.46
LeuL · 6 codons	CUA2.44	CUA2.74	CUA3.52	CUA1.88	CUA2.75	CUA1.73	UUA2.00
IleI · 2 codons	AUC1.00	AUC1.00	AUC1.52	AUU1.40	AUC1.02	AUU1.05	AUU1.46
MetM · 2 codons	AUA1.42	AUA1.58	AUA1.43	AUA1.11	AUA1.62	AUA1.55	AUA1.53
ValV · 4 codons	GUA2.25	GUA2.35	GUA2.45	GUA2.09	GUA1.81	GUA1.97	GUA1.81
SerS · 6 codons	UCA1.82	UCC2.31	UCC1.75	UCA1.75	UCA2.60	UCA1.74	UCA2.00
ProP · 4 codons	CCC1.64	CCC1.60	CCC1.76	CCC1.75	CCA1.65	CCU2.29	CCA2.08
ThrT · 4 codons	ACC1.65	ACA1.74	ACC1.86	ACC1.76	ACC1.71	ACC1.54	ACA1.85
AlaA · 4 codons	GCC1.83	GCA1.70	GCC2.27	GCC1.86	GCC1.91	GCC1.54	GCC1.70
TyrY · 2 codons	UAC1.13	UAU1.03	UAC1.58	UAC1.25	UAC1.21	UAU1.41	UAU1.00
HisH · 2 codons	CAC1.45	CAC1.38	CAC1.64	CAC1.40	CAC1.35	CAU1.03	CAC1.37
GlnQ · 2 codons	CAA1.87	CAA1.93	CAA1.92	CAA1.80	CAA1.86	CAA2.00	CAA2.00
AsnN · 2 codons	AAC1.39	AAC1.24	AAC1.76	AAC1.49	AAC1.48	AAC1.22	AAC1.10
LysK · 2 codons	AAA1.91	AAA1.87	AAA2.00	AAA1.84	AAA1.92	AAA2.00	AAA1.93
AspD · 2 codons	GAC1.45	GAC1.26	GAC1.81	GAC1.48	GAC1.43	GAC1.05	GAU1.02
GluE · 2 codons	GAA1.61	GAA1.63	GAA1.60	GAA1.57	GAA1.74	GAA1.86	GAA1.43
CysC · 2 codons	UGC1.28	UGC1.40	UGC1.43	UGC1.20	UGC1.25	UGU1.18	UGC1.13
TrpW · 2 codons	UGA1.52	UGG2.00	UGA1.74	UGA1.41	UGA1.08	UGA1.80	UGA1.61
ArgR · 4 codons	CGA2.24	CGA3.02	CGA3.08	CGA1.79	CGA2.85	CGC2.41	CGA2.63
GlyG · 4 codons	GGC1.63	GGA1.56	GGC2.26	GGC1.53	GGC1.76	GGC1.56	GGA1.76

RSCU (relative synonymous codon usage) compares a codon with even use of all codons for its amino acid: 1 means no preference, and the maximum equals the number of synonymous codons. Counts are summed over vertebrate species and read with the vertebrate mitochondrial code (NCBI table 2); other animals use different codes and are left out.