YGL186C / TPN1

Plasma membrane pyridoxine (vitamin B6) transporter; member of the purine-cytosine permease subfamily within the major facilitator superfamily; proton symporter with similarity to Fcy21p, Fcy2p, and Fcy22p

Zygosity: Homozygous strain
fixedexpanded
Profile for YGL186C / TPN1

Click on Significant Values for Screen Details ID:SGTC_8|Compound:0354-0001|FD-Score:5.25|P-value:5.37E-4 ID:SGTC_23|Compound:0213-0011|FD-Score:5.71|P-value:1.84E-4 ID:SGTC_61|Compound:0978-8327|FD-Score:7.48|P-value:1.39E-6 ID:SGTC_116|Compound:0155-0182|FD-Score:5.26|P-value:5.32E-4 ID:SGTC_197|Compound:4544-0043|FD-Score:9.42|P-value:1.62E-9 ID:SGTC_301|Compound:4119-0100|FD-Score:7.48|P-value:1.39E-6 ID:SGTC_438|Compound:sphingosine|FD-Score:6.04|P-value:8.10E-5 ID:SGTC_515|Compound:4092-1064|FD-Score:5.73|P-value:1.76E-4 ID:SGTC_526|Compound:4469-0568|FD-Score:8.93|P-value:1.02E-8 ID:SGTC_584|Compound:0986-0246|FD-Score:5.06|P-value:8.18E-4 ID:SGTC_625|Compound:k213-0058|FD-Score:7.84|P-value:4.42E-7 ID:SGTC_712|Compound:r015-0009|FD-Score:7.69|P-value:7.22E-7 ID:SGTC_716|Compound:1683-5115|FD-Score:-6.15|P-value:3.48E-5 ID:SGTC_718|Compound:1130-0073|FD-Score:14.50|P-value:3.68E-20 ID:SGTC_719|Compound:2810-4230|FD-Score:5.26|P-value:5.29E-4 ID:SGTC_720|Compound:r052-1644|FD-Score:-4.96|P-value:6.51E-4 ID:SGTC_729|Compound:4092-0585|FD-Score:-5.49|P-value:1.90E-4 ID:SGTC_788|Compound:1182-0090|FD-Score:6.33|P-value:3.80E-5 ID:SGTC_800|Compound:3448-7362|FD-Score:6.21|P-value:5.26E-5 ID:SGTC_830|Compound:4076-0286|FD-Score:5.81|P-value:1.43E-4 ID:SGTC_860|Compound:0443-0016|FD-Score:5.92|P-value:1.09E-4 ID:SGTC_937|Compound:3122-0005|FD-Score:11.90|P-value:3.54E-14 ID:SGTC_1050|Compound:3232-1156|FD-Score:5.23|P-value:5.61E-4 ID:SGTC_1055|Compound:3448-0672|FD-Score:7.15|P-value:3.79E-6 ID:SGTC_1087|Compound:pterostilbene|FD-Score:5.72|P-value:1.78E-4 ID:SGTC_1130|Compound:3454-2858|FD-Score:9.88|P-value:2.65E-10 ID:SGTC_1159|Compound:4092-0538|FD-Score:5.52|P-value:2.86E-4 ID:SGTC_1165|Compound:k072-0275|FD-Score:5.27|P-value:5.12E-4 ID:SGTC_1260|Compound:0700-1617|FD-Score:-6.07|P-value:4.38E-5 ID:SGTC_1311|Compound:1190-0148|FD-Score:7.40|P-value:1.78E-6 ID:SGTC_1435|Compound:4092-0789|FD-Score:6.27|P-value:4.49E-5 ID:SGTC_1437|Compound:4092-1038|FD-Score:6.21|P-value:5.28E-5 ID:SGTC_1466|Compound:k213-0079|FD-Score:7.41|P-value:1.75E-6 ID:SGTC_1495|Compound:4469-0029|FD-Score:6.11|P-value:6.87E-5 ID:SGTC_1566|Compound:6,7-dimethoxy-4-ethylcoumarin|FD-Score:6.02|P-value:8.47E-5 ID:SGTC_1571|Compound:st070080|FD-Score:-5.37|P-value:2.53E-4 ID:SGTC_1602|Compound:st001878|FD-Score:-8.35|P-value:3.88E-8 ID:SGTC_1606|Compound:st000094|FD-Score:5.09|P-value:7.70E-4 ID:SGTC_1616|Compound:st002734|FD-Score:-6.87|P-value:4.64E-6 ID:SGTC_1638|Compound:st008370|FD-Score:5.05|P-value:8.38E-4 ID:SGTC_1712|Compound:st032288|FD-Score:5.08|P-value:7.80E-4 ID:SGTC_1754|Compound:st042065|FD-Score:7.15|P-value:3.80E-6 ID:SGTC_1788|Compound:fluconazole|FD-Score:5.95|P-value:1.02E-4 ID:SGTC_1826|Compound:st055655|FD-Score:5.06|P-value:8.22E-4 ID:SGTC_1873|Compound:rapamycin|FD-Score:6.62|P-value:1.74E-5 ID:SGTC_1951|Compound:sophoraflavanone G|FD-Score:-4.94|P-value:6.79E-4 ID:SGTC_1969|Compound:st069479|FD-Score:8.14|P-value:1.64E-7 ID:SGTC_1991|Compound:st073000|FD-Score:5.50|P-value:3.01E-4 ID:SGTC_2055|Compound:5235184|FD-Score:6.70|P-value:1.38E-5 ID:SGTC_2093|Compound:5331342|FD-Score:5.24|P-value:5.55E-4 ID:SGTC_2124|Compound:5246320|FD-Score:5.01|P-value:9.19E-4 ID:SGTC_2153|Compound:5569715|FD-Score:5.03|P-value:8.66E-4 ID:SGTC_2177|Compound:5811945|FD-Score:6.59|P-value:1.89E-5 ID:SGTC_2232|Compound:chlorophetanol|FD-Score:7.00|P-value:5.81E-6 ID:SGTC_2240|Compound:6690706|FD-Score:5.93|P-value:1.06E-4 ID:SGTC_2291|Compound:7917025|FD-Score:5.63|P-value:2.21E-4 ID:SGTC_2330|Compound:7979373|FD-Score:10.70|P-value:7.08E-12 ID:SGTC_2332|Compound:7982454|FD-Score:7.40|P-value:1.77E-6 ID:SGTC_2418|Compound:pentamidine|FD-Score:5.63|P-value:2.21E-4 ID:SGTC_2424|Compound:5537928|FD-Score:5.93|P-value:1.07E-4 ID:SGTC_2437|Compound:5741531|FD-Score:-5.71|P-value:1.10E-4 ID:SGTC_2463|Compound:5473429|FD-Score:6.91|P-value:7.75E-6 ID:SGTC_2500|Compound:5-alpha-androstan-3,17-dione|FD-Score:-5.43|P-value:2.17E-4 ID:SGTC_2508|Compound:celastrol|FD-Score:5.19|P-value:6.22E-4 ID:SGTC_2523|Compound:spermine|FD-Score:-6.41|P-value:1.73E-5 ID:SGTC_2525|Compound:harmine|FD-Score:7.25|P-value:2.80E-6 ID:SGTC_2534|Compound:helenine|FD-Score:9.00|P-value:7.85E-9 ID:SGTC_2566|Compound:crystal violet carbinol|FD-Score:5.37|P-value:4.09E-4 ID:SGTC_2568|Compound:diosmetin|FD-Score:-5.54|P-value:1.69E-4 ID:SGTC_2578|Compound:7,8-dihydroxyflavone|FD-Score:-8.64|P-value:1.37E-8 ID:SGTC_2619|Compound:obliquin|FD-Score:5.89|P-value:1.17E-4 ID:SGTC_2623|Compound:humulene|FD-Score:5.98|P-value:9.35E-5 ID:SGTC_2662|Compound:daunorubicin|FD-Score:5.86|P-value:1.29E-4 ID:SGTC_2821|Compound:7996764|FD-Score:9.16|P-value:4.36E-9 ID:SGTC_2906|Compound:7926292|FD-Score:5.77|P-value:1.61E-4 ID:SGTC_2938|Compound:9059746|FD-Score:12.30|P-value:3.77E-15 ID:SGTC_2959|Compound:9079155|FD-Score:8.54|P-value:4.20E-8 ID:SGTC_2963|Compound:9082507|FD-Score:6.21|P-value:5.30E-5 ID:SGTC_3041|Compound:9091929|FD-Score:5.55|P-value:2.71E-4 ID:SGTC_3050|Compound:9093513|FD-Score:7.45|P-value:1.53E-6 ID:SGTC_3054|Compound:9094397|FD-Score:6.64|P-value:1.65E-5 ID:SGTC_3099|Compound:9120367|FD-Score:5.45|P-value:3.42E-4 ID:SGTC_3101|Compound:9120531|FD-Score:5.88|P-value:1.23E-4 ID:SGTC_3138|Compound:9097216|FD-Score:5.46|P-value:3.35E-4 ID:SGTC_3218|Compound:9131285|FD-Score:8.62|P-value:3.09E-8 ID:SGTC_3277|Compound:9138724|FD-Score:7.89|P-value:3.76E-7 ID:SGTC_3286|Compound:9117135|FD-Score:-5.92|P-value:6.37E-5 ID:SGTC_3352|Compound:9151559|FD-Score:5.57|P-value:2.60E-4 ID:SGTC_8|Compound:0354-0001|FD-Score:5.25|P-value:5.37E-4 ID:SGTC_23|Compound:0213-0011|FD-Score:5.71|P-value:1.84E-4 ID:SGTC_61|Compound:0978-8327|FD-Score:7.48|P-value:1.39E-6 ID:SGTC_116|Compound:0155-0182|FD-Score:5.26|P-value:5.32E-4 ID:SGTC_197|Compound:4544-0043|FD-Score:9.42|P-value:1.62E-9 ID:SGTC_301|Compound:4119-0100|FD-Score:7.48|P-value:1.39E-6 ID:SGTC_438|Compound:sphingosine|FD-Score:6.04|P-value:8.10E-5 ID:SGTC_515|Compound:4092-1064|FD-Score:5.73|P-value:1.76E-4 ID:SGTC_526|Compound:4469-0568|FD-Score:8.93|P-value:1.02E-8 ID:SGTC_584|Compound:0986-0246|FD-Score:5.06|P-value:8.18E-4 ID:SGTC_625|Compound:k213-0058|FD-Score:7.84|P-value:4.42E-7 ID:SGTC_712|Compound:r015-0009|FD-Score:7.69|P-value:7.22E-7 ID:SGTC_716|Compound:1683-5115|FD-Score:-6.15|P-value:3.48E-5 ID:SGTC_718|Compound:1130-0073|FD-Score:14.50|P-value:3.68E-20 ID:SGTC_719|Compound:2810-4230|FD-Score:5.26|P-value:5.29E-4 ID:SGTC_720|Compound:r052-1644|FD-Score:-4.96|P-value:6.51E-4 ID:SGTC_729|Compound:4092-0585|FD-Score:-5.49|P-value:1.90E-4 ID:SGTC_788|Compound:1182-0090|FD-Score:6.33|P-value:3.80E-5 ID:SGTC_800|Compound:3448-7362|FD-Score:6.21|P-value:5.26E-5 ID:SGTC_830|Compound:4076-0286|FD-Score:5.81|P-value:1.43E-4 ID:SGTC_860|Compound:0443-0016|FD-Score:5.92|P-value:1.09E-4 ID:SGTC_937|Compound:3122-0005|FD-Score:11.90|P-value:3.54E-14 ID:SGTC_1050|Compound:3232-1156|FD-Score:5.23|P-value:5.61E-4 ID:SGTC_1055|Compound:3448-0672|FD-Score:7.15|P-value:3.79E-6 ID:SGTC_1087|Compound:pterostilbene|FD-Score:5.72|P-value:1.78E-4 ID:SGTC_1130|Compound:3454-2858|FD-Score:9.88|P-value:2.65E-10 ID:SGTC_1159|Compound:4092-0538|FD-Score:5.52|P-value:2.86E-4 ID:SGTC_1165|Compound:k072-0275|FD-Score:5.27|P-value:5.12E-4 ID:SGTC_1260|Compound:0700-1617|FD-Score:-6.07|P-value:4.38E-5 ID:SGTC_1311|Compound:1190-0148|FD-Score:7.40|P-value:1.78E-6 ID:SGTC_1435|Compound:4092-0789|FD-Score:6.27|P-value:4.49E-5 ID:SGTC_1437|Compound:4092-1038|FD-Score:6.21|P-value:5.28E-5 ID:SGTC_1466|Compound:k213-0079|FD-Score:7.41|P-value:1.75E-6 ID:SGTC_1495|Compound:4469-0029|FD-Score:6.11|P-value:6.87E-5 ID:SGTC_1566|Compound:6,7-dimethoxy-4-ethylcoumarin|FD-Score:6.02|P-value:8.47E-5 ID:SGTC_1571|Compound:st070080|FD-Score:-5.37|P-value:2.53E-4 ID:SGTC_1602|Compound:st001878|FD-Score:-8.35|P-value:3.88E-8 ID:SGTC_1606|Compound:st000094|FD-Score:5.09|P-value:7.70E-4 ID:SGTC_1616|Compound:st002734|FD-Score:-6.87|P-value:4.64E-6 ID:SGTC_1638|Compound:st008370|FD-Score:5.05|P-value:8.38E-4 ID:SGTC_1712|Compound:st032288|FD-Score:5.08|P-value:7.80E-4 ID:SGTC_1754|Compound:st042065|FD-Score:7.15|P-value:3.80E-6 ID:SGTC_1788|Compound:fluconazole|FD-Score:5.95|P-value:1.02E-4 ID:SGTC_1826|Compound:st055655|FD-Score:5.06|P-value:8.22E-4 ID:SGTC_1873|Compound:rapamycin|FD-Score:6.62|P-value:1.74E-5 ID:SGTC_1951|Compound:sophoraflavanone G|FD-Score:-4.94|P-value:6.79E-4 ID:SGTC_1969|Compound:st069479|FD-Score:8.14|P-value:1.64E-7 ID:SGTC_1991|Compound:st073000|FD-Score:5.50|P-value:3.01E-4 ID:SGTC_2055|Compound:5235184|FD-Score:6.70|P-value:1.38E-5 ID:SGTC_2093|Compound:5331342|FD-Score:5.24|P-value:5.55E-4 ID:SGTC_2124|Compound:5246320|FD-Score:5.01|P-value:9.19E-4 ID:SGTC_2153|Compound:5569715|FD-Score:5.03|P-value:8.66E-4 ID:SGTC_2177|Compound:5811945|FD-Score:6.59|P-value:1.89E-5 ID:SGTC_2232|Compound:chlorophetanol|FD-Score:7.00|P-value:5.81E-6 ID:SGTC_2240|Compound:6690706|FD-Score:5.93|P-value:1.06E-4 ID:SGTC_2291|Compound:7917025|FD-Score:5.63|P-value:2.21E-4 ID:SGTC_2330|Compound:7979373|FD-Score:10.70|P-value:7.08E-12 ID:SGTC_2332|Compound:7982454|FD-Score:7.40|P-value:1.77E-6 ID:SGTC_2418|Compound:pentamidine|FD-Score:5.63|P-value:2.21E-4 ID:SGTC_2424|Compound:5537928|FD-Score:5.93|P-value:1.07E-4 ID:SGTC_2437|Compound:5741531|FD-Score:-5.71|P-value:1.10E-4 ID:SGTC_2463|Compound:5473429|FD-Score:6.91|P-value:7.75E-6 ID:SGTC_2500|Compound:5-alpha-androstan-3,17-dione|FD-Score:-5.43|P-value:2.17E-4 ID:SGTC_2508|Compound:celastrol|FD-Score:5.19|P-value:6.22E-4 ID:SGTC_2523|Compound:spermine|FD-Score:-6.41|P-value:1.73E-5 ID:SGTC_2525|Compound:harmine|FD-Score:7.25|P-value:2.80E-6 ID:SGTC_2534|Compound:helenine|FD-Score:9.00|P-value:7.85E-9 ID:SGTC_2566|Compound:crystal violet carbinol|FD-Score:5.37|P-value:4.09E-4 ID:SGTC_2568|Compound:diosmetin|FD-Score:-5.54|P-value:1.69E-4 ID:SGTC_2578|Compound:7,8-dihydroxyflavone|FD-Score:-8.64|P-value:1.37E-8 ID:SGTC_2619|Compound:obliquin|FD-Score:5.89|P-value:1.17E-4 ID:SGTC_2623|Compound:humulene|FD-Score:5.98|P-value:9.35E-5 ID:SGTC_2662|Compound:daunorubicin|FD-Score:5.86|P-value:1.29E-4 ID:SGTC_2821|Compound:7996764|FD-Score:9.16|P-value:4.36E-9 ID:SGTC_2906|Compound:7926292|FD-Score:5.77|P-value:1.61E-4 ID:SGTC_2938|Compound:9059746|FD-Score:12.30|P-value:3.77E-15 ID:SGTC_2959|Compound:9079155|FD-Score:8.54|P-value:4.20E-8 ID:SGTC_2963|Compound:9082507|FD-Score:6.21|P-value:5.30E-5 ID:SGTC_3041|Compound:9091929|FD-Score:5.55|P-value:2.71E-4 ID:SGTC_3050|Compound:9093513|FD-Score:7.45|P-value:1.53E-6 ID:SGTC_3054|Compound:9094397|FD-Score:6.64|P-value:1.65E-5 ID:SGTC_3099|Compound:9120367|FD-Score:5.45|P-value:3.42E-4 ID:SGTC_3101|Compound:9120531|FD-Score:5.88|P-value:1.23E-4 ID:SGTC_3138|Compound:9097216|FD-Score:5.46|P-value:3.35E-4 ID:SGTC_3218|Compound:9131285|FD-Score:8.62|P-value:3.09E-8 ID:SGTC_3277|Compound:9138724|FD-Score:7.89|P-value:3.76E-7 ID:SGTC_3286|Compound:9117135|FD-Score:-5.92|P-value:6.37E-5 ID:SGTC_3352|Compound:9151559|FD-Score:5.57|P-value:2.60E-4

Top fitness defect scores for YGL186C deletion by condition

RankScreen IDConditionFD ScoreConc
P-value
Response
Signature
Screen
Rank
Screen
P-value
1 SGTC_718 1130-0073 14.50 349.0 µM 3.68E-20 1 1.08E-47
2 SGTC_2938 9059746 12.30 27.3 µM 3.77E-15 fatty acid desaturase (OLE1) 3 2.76E-35
3 SGTC_937 3122-0005 11.90 46.7 µM 3.54E-14 1 7.03E-33
4 SGTC_2330 7979373 10.70 200.0 µM 7.08E-12 60S ribosome export 2 3.41E-27
5 SGTC_1130 3454-2858 9.88 15.9 µM 2.65E-10 NEO1 16 2.60E-23
6 SGTC_197 4544-0043 9.42 41.6 µM 1.62E-9 endomembrane recycling 6 2.26E-21
7 SGTC_2821 7996764 9.16 71.4 µM 4.36E-9 1 2.60E-20
8 SGTC_2534 helenine 9.00 20.1 µM 7.85E-9 RSC complex & mRNA processing 2 1.11E-19
9 SGTC_526 4469-0568 8.93 170.0 µM 1.02E-8 azole & statin 7 2.13E-19
10 SGTC_3218 9131285 8.62 49.5 µM 3.09E-8 endomembrane recycling 6 3.23E-18
11 SGTC_2959 9079155 8.54 11.0 µM 4.20E-8 2 6.87E-18
12 SGTC_1969 st069479 8.14 40.1 µM 1.64E-7 1 1.97E-16
13 SGTC_3277 9138724 7.89 49.5 µM 3.76E-7 Golgi 5 1.50E-15
14 SGTC_625 k213-0058 7.84 121.0 µM 4.42E-7 translation 12 2.24E-15
15 SGTC_712 r015-0009 7.69 80.6 µM 7.22E-7 4 7.48E-15
16 SGTC_61 0978-8327 7.48 98.6 µM 1.39E-6 ergosterol biosynthesis 1 3.70E-14
17 SGTC_301 4119-0100 7.48 24.5 µM 1.39E-6 NEO1 24 3.70E-14
18 SGTC_3050 9093513 7.45 49.5 µM 1.53E-6 3 4.73E-14
19 SGTC_1466 k213-0079 7.41 57.5 µM 1.75E-6 2 6.56E-14
20 SGTC_2332 7982454 7.40 138.9 µM 1.77E-6 6 6.75E-14

Download Fitness data (tab-delimited text)  (excel)
Cofit Genes
Download Cofitness data (tab-delimited text)  (excel)

Correlation pval ORF Gene Zygosity Description
0.165 6.18E-22 YDL118W_d YDL118W_d hom Dubious open reading frame, unlikely to encode a protein; overlaps almost completely with YDL119C; mutations that would affect both YDL118W and YDL119C confer defective telomere maintenance and are synthetically sick or lethal with alpha-synuclein
0.134 5.89E-15 YMR202W ERG2 hom C-8 sterol isomerase, catalyzes the isomerization of the delta-8 double bond to the delta-7 position at an intermediate step in ergosterol biosynthesis
0.127 1.64E-13 YDL119C_p YDL119C_p hom Putative mitochondrial transport protein; GFP-fusion protein is induced in response to the DNA-damaging agent MMS; the authentic, non-tagged protein is detected in purified mitochondria
0.124 6.09E-13 YEL029C BUD16 hom Putative pyridoxal kinase, a key enzyme involved in pyridoxal 5'-phosphate synthesis, the active form of vitamin B6; required for genome integrity; involved in bud-site selection; similarity to yeast BUD17 and human pyridoxal kinase (PDXK)
0.118 7.11E-12 YMR154C RIM13 hom Calpain-like cysteine protease; involved in proteolytic activation of Rim101p in response to alkaline pH; localizes to punctate structures in alkaline conditions and in vps4 mutant; has similarity to A. nidulans palB
0.117 9.33E-12 YJL204C RCY1 hom F-box protein involved in recycling plasma membrane proteins internalized by endocytosis; localized to sites of polarized growth
0.117 1.33E-11 YOR293W RPS10A hom Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S10, no bacterial homolog; RPS10A has a paralog, RPS10B, that arose from the whole genome duplication
0.116 1.46E-11 YCR094W CDC50 hom Endosomal protein that interacts with phospholipid flippase Drs2p; interaction with Cdc50p is essential for Drs2p catalytic activity; mutations affect cell polarity and polarized growth; similar to Lem3p; CDC50 has a paralog, YNR048W, that arose from the whole genome duplication
0.116 1.69E-11 YKL212W SAC1 hom Phosphatidylinositol phosphate (PtdInsP) phosphatase; involved in hydrolysis of PtdIns[4]P in the early and medial Golgi; regulated by interaction with Vps74p; ER localized transmembrane protein which cycles through the Golgi; involved in protein trafficking and processing, secretion, and cell wall maintenance; regulates sphingolipid biosynthesis through the modulation of PtdIns(4)P metabolism
0.115 2.45E-11 YGR122W YGR122W hom Protein that may be involved in pH regulation; probable ortholog of A. nidulans PalC, which is involved in pH regulation and binds to the ESCRT-III complex; null mutant does not properly process Rim101p and has decreased resistance to rapamycin; GFP-fusion protein is cytoplasmic; relative distribution to cytoplasm increases upon DNA replication stress
0.115 2.94E-11 YGL054C ERV14 hom COPII-coated vesicle protein; involved in vesicle formation and incorporation of specific secretory cargo; required for the delivery of bud-site selection protein Axl2p to cell surface; related to Drosophila cornichon; ERV14 has a paralog, ERV15, that arose from the whole genome duplication
0.114 4.25E-11 YLR025W SNF7 hom One of four subunits of the endosomal sorting complex required for transport III (ESCRT-III); involved in the sorting of transmembrane proteins into the multivesicular body (MVB) pathway; recruited from the cytoplasm to endosomal membranes
0.113 5.22E-11 YOR070C GYP1 hom Cis-golgi GTPase-activating protein (GAP) for the Rab family members Ypt1p (in vivo) and for Ypt1p, Sec4p, Ypt7p, and Ypt51p (in vitro); involved in vesicle docking and fusion
0.111 1.09E-10 YMR063W RIM9 hom Plasma membrane protein of unknown function; involved in the proteolytic activation of Rim101p in response to alkaline pH; interacts with Rim21p and Dfg16p to form a pH-sensing complex in the Rim101 pathway and is required to maintain Rim21p levels; has similarity to A. nidulans PalI;
0.105 9.53E-10 YMR077C VPS20 hom Myristoylated subunit of ESCRTIII, the endosomal sorting complex required for transport of transmembrane proteins into the multivesicular body pathway to the lysosomal/vacuolar lumen; cytoplasmic protein recruited to endosomal membranes