Biomineraliztion in dark matter

where in the WGCBNA and barnacle?
e5
barnacle
coral
Author
Affiliation

Steven Roberts

Published

November 15, 2025

Have list of biomineralization genes from coral literature (e.g., Takeuchi et al. 2016, Ramos-Silva et al. 2013)

Accession / GeneID Blasted protein in Stylophora Definition Reference
AAD11470.1 XP_022803851.1 L-type calcium channel alpha-1 subunit Zoccola et al., 1999
AAR13013.1 XP_022785723.1 plasma membrane calcium ATPase Zoccola et al., 2004
ACE95141.1 XP_022799914.1 carbonic anhydrase Moya et al., 2008
AGE35225.2 XP_022785805.1 CARP1 Mass et al., 2013
AGE35226.1 XP_022794102.1 CARP3 Mass et al., 2013
AGG36361.1 XP_022784808.1 Protoacadherin (PC4) Drake et al., 2013
AJQ31790.1 XP_022788307.1 solute carrier family 4 member gamma Zoccola et al., 2015
aug_v2a.00002.t1 XP_022808048.1, XP_022808045.1, XP_022790310.1 EP-like 2 Takeuchi et al., 2016
aug_v2a.01440.t1 (aug_v2a.01441.t1) XP_022780690.1 Adi-SAARP2 Takeuchi et al., 2016
aug_v2a.02830 XP_022782704.1, XP_022782703.1 PKD1-related protein Takeuchi et al., 2016
aug_v2a.02830.t1 XP_022782704.1, XP_022782703.1 PKD1-related protein Takeuchi et al., 2016
aug_v2a.05945.t1 XP_022783430.1 TSP-1 and VWA domain-containing Takeuchi et al., 2016
aug_v2a.06122.t1 XP_022804012.1 EGF & laminin G domain protein Takeuchi et al., 2016
aug_v2a.06123.t1 XP_022804012.1 EGF & laminin G domain protein Takeuchi et al., 2016
aug_v2a.06327.t1 XP_022781839.1, XP_022781915.1 SAARP3 Takeuchi et al., 2016
aug_v2a.07627.t1 XP_022806326.1 Zona pellucida domain protein Takeuchi et al., 2016
aug_v2a.09809.t1 XP_022801334.1 Mucin4-like protein Takeuchi et al., 2016
aug_v2a.09968.t1 XP_022794736.1 MAM & LDLr domain protein Takeuchi et al., 2016
aug_v2a.09969.t1 XP_022794736.1 MAM & LDLr domain protein Takeuchi et al., 2016
aug_v2a.11068.t1 XP_022782398.1 SAARP1 Takeuchi et al., 2016
aug_v2a.15064.t1 XP_022808152.1 Cysteine-rich Takeuchi et al., 2016
aug_v2a.15065.t1 XP_022794122.1, XP_022794121.1 galaxin2 Takeuchi et al., 2016
aug_v2a.15580.t1 XP_022804012.1 Laminin G domain protein Takeuchi et al., 2016
aug_v2a.18631.t1 XP_022794122.1, XP_022794121.1 galaxin Takeuchi et al., 2016
aug_v2a.19518.t1 XP_022784808.1 Protocadherin-like Takeuchi et al., 2016
aug_v2a.22918.t1 XP_022808048.1, XP_022808045.1, XP_022790310.1 EP-like1 Takeuchi et al., 2016
aug_v2a.24015.t1 XP_022788227.1 Hephaestin-like protein Takeuchi et al., 2016
aug_v2a.24512.t1 XP_022804012.1 EGF & laminin G domain protein Takeuchi et al., 2016
EU532164.1 XP_022799914.1 carbonic anhydrase 2 Bertucci et al., 2011
Gene:g29033.t1 XP_022799914.1 Carbonic Anhydrase (STPCA2-1) Mummadisetti et al., 2021
Gene:g3745.t1 XP_022782398.1 CARP4 Mummadisetti et al., 2021
Gene:10186 XP_022784808.1 Protocadherin (PC2) Mummadisetti et al., 2021
Gene:g10186 XP_022782398.1 CARP4 Mummadisetti et al., 2021
Gene:g10187 XP_022784808.1 Protocadherin (PC3) Mummadisetti et al., 2021
Gene:g10188 XP_022784808.1 Protocadherin (PC3) Mummadisetti et al., 2021
Gene:g11190 XP_022783323.1 USOMP12 Mummadisetti et al., 2021
Gene:g13552 XP_022781839.1 Acidic SOMP (p27) Mummadisetti et al., 2021
Gene:g14733 XP_022782398.1 CARP4 Mummadisetti et al., 2021
Gene:g1484 XP_022785805.1 CARP1 Mummadisetti et al., 2021
Gene:g15294.t1 XP_022779720.1 Vitellogenin Mummadisetti et al., 2021
Gene:g15955 XP_022794736.1 MAM LDL-2 Mummadisetti et al., 2021
Gene:g2115 XP_022784808.1 Cadherin Mummadisetti et al., 2021
Gene:g2116 XP_022784808.1 Protocadherin (PC1) Mummadisetti et al., 2021
Gene:g22569 XP_022810031.1 Fibronectin Mummadisetti et al., 2021
Gene:g24177 XP_022798258.1 Protocadherin (PC5) Mummadisetti et al., 2021
Gene:g27814 XP_022801359.1 carbonic anhydrase (STPCA2-2) Mummadisetti et al., 2021
Gene:g2829 XP_022783414.1, XP_022783413.1 Thrombospondin Mummadisetti et al., 2021
Gene:g2829.t1 XP_022783415.1 Coadhesin Mummadisetti et al., 2021
Gene:g29034.t1 XP_022799914.1 Carbonic Anhydrase Mummadisetti et al., 2021
Gene:g30 XP_022784808.1 Protoacadherin (PC4) Mummadisetti et al., 2021
Gene:g30385.t1 XP_022780049.1 USOMPS13 Mummadisetti et al., 2021
Gene:g34749 XP_022804012.1 EGF & LamininG-Like Mummadisetti et al., 2021
Gene:g37058 XP_022804608.1 Fibronectin-2 Mummadisetti et al., 2021
Gene:g38128 XP_022783430.1 α-Collagen Mummadisetti et al., 2021
Gene:g38881 XP_022790556.1, XP_022790555.1 USOMP14 Mummadisetti et al., 2021
Gene:g39770 XP_022795871.1 Kielin-Like Mummadisetti et al., 2021
Gene:g5735.t1 XP_022799127.1 Tolloid-Like Mummadisetti et al., 2021
Gene:g7086 XP_022804012.1 EGF LamG1 Mummadisetti et al., 2021
Gene:g8396 XP_022803859.1 CARP6-partial Mummadisetti et al., 2021
Gene:g907 XP_022806326.1 Zona Pellucida Mummadisetti et al., 2021
Gene:g9094 XP_022795459.1, XP_022777933.1 Actin Mummadisetti et al., 2021
HM163215 XP_029187982.1 Acropora millepora galaxin Reyes-Bermudez et al., 2009
JN631095.1 XP_029195765.1 hypothetical protein p251_4 Hayward et al., 2011
JR971508.1 XP_022793420.1, XP_022793422.1 USOMP6 Ramos-Silva et al., 2013
JR972076.1 XP_022781839.1, XP_022781915.1 Acidic SOMP Ramos-Silva et al., 2013
JR973117.1 XP_022808048.1, XP_022808045.1 USOMP-5 Ramos-Silva et al., 2013
JR976690.1 XP_022794122.1, XP_022794121.1 Galaxin 2 Ramos-Silva et al., 2013
JR980881.1 XP_022804012.1 EGF & laminin G protein Ramos-Silva et al., 2013
JR982706.1 XP_022808152.1 USOMP-2 Ramos-Silva et al., 2013
JR986059.1 XP_022782738.1 Cephalotoxin-like Ramos-Silva et al., 2013
JR991083.1 XP_022779335.1 Collagen α-1 chain Ramos-Silva et al., 2013
JR991407.1 XP_022780690.1 SAARP2 Ramos-Silva et al., 2013
JR994474.1 XP_022794736.1 MAM & LDL-receptor protein 2 Ramos-Silva et al., 2013
JR997000.1 XP_022807628.1, XP_022807627.1 USOMP-3 Ramos-Silva et al., 2013
JR998014.1 XP_022799897.1, XP_022801331.1 Putative carbonic anhydrase Ramos-Silva et al., 2013
JR998260.1 XP_022803212.1 USOMP7 Ramos-Silva et al., 2013
JT001945.1 XP_022782398.1 SAARP1 Ramos-Silva et al., 2013
JT004498.1 XP_022780690.1 USOMP-4 Ramos-Silva et al., 2013
JT011118.1 XP_022794736.1 MAM LDL receptor protein 1 Ramos-Silva et al., 2013
JT014391.1 XP_022797021.1 USOMP-8 Ramos-Silva et al., 2013
JT016638.1 XP_022783430.1 Coadhesin Ramos-Silva et al., 2013
MG182344.1 XP_022792788.1 Acropora yongei Na+/Ca2+ exchanger Barron et al., 2018
MG182345.1 XP_022784107.1 Acropora yongei Na+/Ca2+ exchanger Barron et al., 2018
P12_g2385 XP_022796981.1 hypothetical protein Drake et al., 2013
P13_g6918 XP_022803207.1 Sushi domain-containing Drake et al., 2013
P14_g9951 XP_022783415.1 collagen-like protein Drake et al., 2013
P15_g1532 XP_022780640.1, XP_022783414.1 CARP9 Drake et al., 2013
P16_g11702 XP_022780049.1 hypothetical protein Drake et al., 2013
P18_g810 XP_022783430.1 collagen-like protein Drake et al., 2013
P19_g20041 XP_022804012.1 Contactin-associated protein Drake et al., 2013
P2_g11187 XP_022782364.1 CARP8 Drake et al., 2013
P20_g6066 XP_022794736.1 MAM domain anchor protein Drake et al., 2013
P21_g18277 XP_022806326.1 Zona pellucida Drake et al., 2013
P22_g19762 XP_022785540.1 hypothetical protein Drake et al., 2013
P23_g1057 XP_022808083.1, XP_022808082.1 Protocadherin Drake et al., 2013
P24_g15888 XP_022779720.1 vitellogenin-like protein Drake et al., 2013
P26_g1441 XP_022794920.1 vitellogenin-like protein Drake et al., 2013
P27_g18472 XP_022781915.1 Integrin-alpha Drake et al., 2013
P28_g11651 XP_022793421.1 Late embryogenesis protein Drake et al., 2013
P3_g12510 XP_022783415.1 Thrombospondin Drake et al., 2013
P31_g20420 XP_022804012.1 Neurexin Drake et al., 2013
P32_g5540 XP_022795871.1 Kielin/chordin-like Drake et al., 2013
P33_g8985 XP_022781364.1 Flagellar associated protein Drake et al., 2013
P34_g1714 XP_022794736.1 MAM/LDL receptor domain protein Drake et al., 2013
P36_g13890 XP_022794736.1 Zonadhesion-like precursor Drake et al., 2013
P4_g9861 XP_022797264.1 Viral inclusion protein Drake et al., 2013
P5_g11674 XP_022783415.1 Hemicentin Drake et al., 2013
P8_g9654 XP_022786917.1, XP_022786918.1 Major yolk protein Drake et al., 2013
P9_g10811; P1_g11108; P10_g11107 XP_022784808.1 Protocadherin fat-like Drake et al., 2013
PFX13778.1 XP_022808646.1 Sacsin Peled et al., 2020
PFX14205.1 XP_022808089.1 Ret receptor kinase (partial) Peled et al., 2020
PFX15740.1 XP_022806058.1 Protein FAM208A Peled et al., 2020
PFX16398.1 XP_022805070.1 hypothetical protein Peled et al., 2020
PFX18785.1 XP_022801334.1, XP_022801335.1 Mucin-4 Peled et al., 2020
PFX26597.1 XP_022788977.1 Complement C3 Peled et al., 2020
PFX26751.1 XP_022788730.1 Transmembrane protease serine 9 Peled et al., 2020
PFX27832.1 XP_022786864.1 PARP11 Peled et al., 2020
PFX30903.1 XP_022781839.1 hypothetical protein Peled et al., 2020
XP_022778254.1 XP_022778254.1 uncharacterized protein LOC111319781 Peled et al., 2020
XP_022778283.1 XP_022778283.1 uncharacterized protein LOC111319816 Peled et al., 2020
XP_022779720.1 XP_022779720.1 vitellogenin-like Peled et al., 2020
XP_022780303.1 XP_022780303.1 uncharacterized protein LOC111321626 Peled et al., 2020
XP_022780690.1 XP_022780690.1 CARP5 Peled et al., 2020
XP_022780694.1 XP_022780694.1 CUB & peptidase domain protein Peled et al., 2020
XP_022781731.1 XP_022781731.1 sulfate anion transporter-like 2C Zoccola et al., 2015
XP_022782398.1 XP_022782398.1 CARP4 Peled et al., 2020
XP_022783031.1 XP_022783031.1 sodium bicarbonate transporter-like 11 Zoccola et al., 2015
XP_022783044.1 XP_022783044.1 uncharacterized protein LOC111323869 Peled et al., 2020
XP_022783323.1 XP_022783323.1 VHS3-like Peled et al., 2020
XP_022783415.1 XP_022783415.1 coadhesin-like isoform X3 Peled et al., 2020
XP_022783952.1 XP_022783952.1 collagenase 3-like Peled et al., 2020
XP_022784623.1 XP_022784623.1 sperm-associated cation channel Peled et al., 2020
XP_022786582.1 XP_022786582.1 synapsin-2-like isoform X2 Peled et al., 2020
XP_022786918.1 XP_022786918.1 major yolk protein-like X2 Peled et al., 2020
XP_022788227.1 XP_022788227.1 hephaestin-like protein Peled et al., 2020
XP_022788270.1 XP_022788270.1 band 3 anion transport protein-like Zoccola et al., 2015
XP_022788730.1 XP_022788730.1 elastase-like peptidase Peled et al., 2020
XP_022789591.1 XP_022789591.1 endothelin-converting enzyme 1-like X2 Peled et al., 2020
XP_022789932.1 XP_022789932.1 MAGUK p55 subfamily member 7-like Peled et al., 2020
XP_022790441.1 XP_022790441.1 PHD finger protein 21A-like Peled et al., 2020
XP_022791567.1 XP_022791567.1 anion exchange protein 2-like Zoccola et al., 2015
XP_022792212.1 XP_022792212.1 ras-like protein 3 Peled et al., 2020
XP_022794122.1 XP_022794122.1 galaxin-like isoform X2 Peled et al., 2020
XP_022794351.1 XP_022794351.1 ependymin-related protein Peled et al., 2020
XP_022794736.1 XP_022794736.1 MAM & LDL receptor protein 2-like Peled et al., 2020
XP_022796981.1 XP_022796981.1 skeletal organic matrix protein 8-like Peled et al., 2020
XP_022796982.1 XP_022796982.1 uncharacterized protein LOC111335364 Peled et al., 2020
XP_022798902.1 XP_022798902.1 LDL receptor-related protein 8-like Peled et al., 2020
XP_022799089.1 XP_022799089.1 CUB domain protein isoform X2 Peled et al., 2020
XP_022799541.1 XP_022799541.1 uncharacterized protein LOC111337489 Peled et al., 2020
XP_022800320.1 XP_022800320.1 sulfate anion transporter 1-like X1 Zoccola et al., 2015
XP_022800329.1 XP_022800329.1 sulfate anion transporter 1-like X2 Zoccola et al., 2015
XP_022800339.1 XP_022800339.1 sulfate anion transporter 1-like X1 Zoccola et al., 2015
XP_022801463.1 XP_022801463.1 bicarbonate cotransporter 3-like X2 Zoccola et al., 2015
XP_022803524.1 XP_022803524.1 digestive cysteine proteinase 1-like Peled et al., 2020
XP_022803808.1 XP_022803808.1 tumor suppressor DMBT1-like Peled et al., 2020
XP_022803872.1 XP_022803872.1 spore wall protein 2-like X3 Peled et al., 2020
XP_022803894.1 XP_022803894.1 uncharacterized protein LOC111341206 Peled et al., 2020
XP_022804012.1 XP_022804012.1 EGF & laminin G domain protein Peled et al., 2020
XP_022804785.1 XP_022804785.1 thioredoxin reductase 1-like Peled et al., 2020
XP_022805470.1 XP_022805470.1 uncharacterized protein LOC111342641 Peled et al., 2020
XP_022806326.1 XP_022806326.1 ZP domain protein Peled et al., 2020
XP_022806664.1 XP_022806664.1 protein lingerer-like Peled et al., 2020
XP_022806928.1 XP_022806928.1 SLIT-ROBO RhoGAP1-like Peled et al., 2020
XP_022807143.1 XP_022807143.1 condensin-2 subunit D3-like Peled et al., 2020
XP_022807256.1 XP_022807256.1 uncharacterized protein LOC111344300 Peled et al., 2020
XP_022807807.1 XP_022807807.1 uncharacterized protein LOC111344812 Peled et al., 2020
XP_022808163.1 XP_022808163.1 uncharacterized protein LOC111345150 Peled et al., 2020
XP_022808576.1 XP_022808576.1 LOC111345553 isoform X2 Peled et al., 2020
XP_022809269.1 XP_022809269.1 microtubule-associated tumor suppressor 1-like X1 Peled et al., 2020
XP_022809270.1 XP_022809270.1 microtubule-associated tumor suppressor 1-like X2 Peled et al., 2020
XP_022810585.1 XP_022810585.1 vWF D + EGF domain protein (partial) Peled et al., 2020
seqkit grep -f ../data/biomin-ids.txt ../data/GCF_002571385.2_Stylophora_pistillata_v1.1_protein.faa > ../output/31-biomin-pathway/Stylo-biomin.fasta
blastp \
  -query ../output/11.3-ortholog-annotation/representative_sequences.faa \
  -db ../output/31-biomin-pathway/Stylo-biomin-db \
  -out ../output/31-biomin-pathway/OG-repseq_Stylo-biomin-results.tsv \
  -outfmt "6 qseqid sseqid evalue bitscore" \
  -evalue 1e-20 \
  -max_target_seqs 1 \
  -max_hsps 1 \
  -num_threads 10
head ../output/31-biomin-pathway/OG-repseq_Stylo-biomin-results.tsv
FUN_000236-T1   XP_022780303.1  6.14e-24    102
FUN_000857-T1   XP_022795459.1  1.39e-159   447
FUN_000867-T1   XP_022780694.1  1.60e-34    125
FUN_001009-T1   XP_022789591.1  7.04e-107   338
FUN_001088-T1   XP_022783044.1  3.90e-35    130
FUN_001089-T1   XP_022783044.1  2.17e-23    93.2
FUN_001101-T1   XP_022783044.1  1.22e-31    118
FUN_001107-T1   XP_022795459.1  9.45e-59    190
FUN_001118-T1   XP_022801359.1  1.33e-35    124
FUN_001185-T1   XP_022798902.1  0.0 870
head ../output/11.3-ortholog-annotation/gene_to_ortholog_mapping.csv
gene_id,species,group_id,type,conservation_level
FUN_000185-T1,Apul,OG_00001,three_way,medium
Peve_00037402,Peve,OG_00001,three_way,medium
Pocillopora_meandrina_HIv1___RNAseq.g28886.t1,Ptua,OG_00001,three_way,medium
FUN_000189-T1,Apul,OG_00002,three_way,medium
Peve_00038462,Peve,OG_00002,three_way,medium
Pocillopora_meandrina_HIv1___RNAseq.g28888.t3,Ptua,OG_00002,three_way,medium
FUN_000190-T1,Apul,OG_00003,three_way,low
Peve_00038463,Peve,OG_00003,three_way,low
Pocillopora_meandrina_HIv1___RNAseq.g28889.t1,Ptua,OG_00003,three_way,low
library(dplyr)
library(readr)

# Paths
blast_file <- "../output/31-biomin-pathway/OG-repseq_Stylo-biomin-results.tsv"
mapping_file <- "../output/11.3-ortholog-annotation/gene_to_ortholog_mapping.csv"

# Read BLAST results
blast <- read_tsv(
  blast_file,
  col_names = c("gene_id", "hit", "evalue", "score")
)

# Read ortholog mapping
mapping <- read_csv(mapping_file)

# Join to add group_id to each gene in blast results
annotated <- blast %>%
  left_join(mapping %>% select(gene_id, group_id), by = "gene_id")


# Write out CSV
write_csv(annotated, "../output/31-biomin-pathway/OG-repseq_Stylo-biomin.csv")


# View result
annotated

Matches in Barnacle

library(tidyverse)

# 1. Load the Stylo-biomin group_id list
stylo_file <- "../output/31-biomin-pathway/OG-repseq_Stylo-biomin.csv"

stylo_groups <- read_csv(stylo_file) %>%
  pull(group_id) %>%
  unique()

# 2. Get all *_top100.csv files
path_top100 <- "../output/22-Visualizing-Rank-outs"

top100_files <- list.files(
  path_top100,
  pattern = "*_top100.csv",
  full.names = TRUE
)

# 3. For each file, find matching group IDs in column 1
results <- map_df(top100_files, function(f) {

  df <- read_csv(f, col_names = FALSE)

  # assume the first column has the group_ids
  col1_ids <- df[[1]] %>% unique()

  matches <- intersect(stylo_groups, col1_ids)

  tibble(
    file = basename(f),
    matched_group_id = matches
  )
})

# Show results
results

# 4. Write out summary file
write_csv(
  results,
  "../output/31-biomin-pathway/Stylo-biomin_matches_in_top100.csv"
)
library(tidyverse)

# Read the file: it has two columns (file, group_id)
res <- read_csv("../output/31-biomin-pathway/Stylo-biomin_matches_in_top100.csv",
                col_names = c("file", "group_id"))

# Count occurrences of each file (column 1)
file_counts <- res %>%
  count(file, name = "n_occurrences") %>%
  arrange(desc(n_occurrences))

file_counts

WGCNA module enrichment

library(tidyverse)

# 1. Load biomin group IDs
biomin_groups <- read_csv("../output/31-biomin-pathway/OG-repseq_Stylo-biomin.csv") %>%
  pull(group_id) %>%
  unique()

n_biomin <- length(biomin_groups)  # total number of biomin groups

# 2. Load module assignments (must contain group_id and module_label)
assignments <- read_csv(
  "~/GitHub/timeseries_molecular/M-multi-species/output/18-ortholog-wgcna/gene_module_assignments.csv"
)

# 3. Mark biomin matches
assignments2 <- assignments %>%
  mutate(is_biomin = group_id %in% biomin_groups)

# 4. Summarize by module_label
module_summary <- assignments2 %>%
  group_by(module_label) %>%
  summarise(
    n_group_ids_in_module = n_distinct(group_id),
    n_biomin_in_module = sum(is_biomin),
    
    # percent of BIOMIN groups captured by this module
    pct_of_all_biomin = (n_biomin_in_module / n_biomin) * 100,
    
    # percent of module that is biomin
    pct_of_module_that_is_biomin = (n_biomin_in_module / n_group_ids_in_module) * 100
  ) %>%
  arrange(desc(pct_of_all_biomin))

module_summary