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To determine the influence of the β-secretase cleavage pathway on the occurrence of endogenous nuclear AICD spots, primary neurons were treated with pharmacological inhibitors to modulate cleavage of either α- or β-secretase. In order to test the efficiency of commercially available inhibitors, neurons were treated for 22 hours at DIV 14 with 20 μM of the α-secretase inhibitor TAPI 1, or with 60 nM BACE1 inhibitor A, or with 1.4 μM BACE1 inhibitor B. In addition, cells were treated concomitantly with 1 μM γ-secretase inhibitor DAPT to accumulate CTFs. As expected, treatment with TAPI 1 resulted in a reduction in α-CTF levels, whereas treatment with either BACE1 inhibitor A or B resulted in a reduction in β-CTFs, when compared to treatment with DAPT alone ( Fig. 8C ). Neurons were subsequently treated with the different secretase inhibitors together with 20 ng/ml LMB for 22 hours, followed by staining with APP C-terminal and Map2ab antibodies, and analysis by confocal microscopy ( Fig. 8D ).

Neurons were identified by the Map2ab staining that also guided the identification of the optimal mid-nuclear plane as described above. Single confocal sections from 19-42 neurons were imaged for each condition (control, n =19; LMB, n =35; DAPT + LMB, n =34; TAPI 1 + LMB, n =42; BACE1 inhibitor A + LMB, n =38; BACE1 inhibitor B + LMB, n =22). The nuclei were outlined with the help of the Map2ab staining, and the number of AICD spots quantified by two independent researchers blinded to the experimental conditions ( Fig. 8E ). After treatment with LMB an increased number of AICD spots was clearly visible in the nucleus, which was not significantly reduced by TAPI 1 co-treatment. By contrast, treatment of neurons with DAPT, which prevents AICD generation, or with either of the BACE1 inhibitors resulted in a significant inhibition of nuclear AICD spot formation (MWU, P <0.001 versus LMB alone). Thus, the inhibition of β-secretase was sufficient to prevent nuclear signaling of endogenous AICD in neurons, whereas α-secretase inhibition had no effect.

Fig. 8.

BACE1 inhibition in primary neurons inhibits endogenous AICD nuclear signaling. (A) Primary neuronal cultures, prepared from postnatal day zero C57/Bl6 mice, were treated at DIV 14 with 20 ng/ml LMB, followed by nuclei isolation and immunoprecipitation. Western blotting with anti-APP C-terminal antibody shows a dramatic increase in the level of AICD detected in the nuclear fraction after LMB treatment. (B) Neurons treated with LMB were fixed in methanol and stained for APP, Map2ab and DNA (DRAQ5). Confocal sections were taken 1 μm apart in the -axis. The first panel shows the mid-nuclear plane, free of Map2ab staining in the DRAQ5-stained nuclei (insert). Moving towards the bottom of the cell the second panel still seems to encompass the nucleus according to the DRAQ5 staining but the Map2ab staining shows that microtubules already overlay with the nuclear staining. The third panel shows the microtubule network lying beneath the nucleus with APP staining in vesicular structures. Scale bar: 7 μm. (C) Primary neuronal cultures at 14 DIV were treated with 20 μM α-secretase inhibitor TAPI 1, 60 nM BACE1 inhibitor A (Calbiochem, #171601) or 1.4 μM BACE1 inhibitor B (Calbiochem, #565749), together with 1 μM γ-secretase inhibitor DAPT to accumulate CTFs. Western blots were stained with APP C-terminal antibodies. Treatment with TAPI 1 leads to a reduction in levels of α-CTFs, whereas treatment with either BACE1 inhibitor A or B leads to a reduction in β-CTFs, when compared to DAPT treatment alone. (D) Neuronal cultures at DIV 14 were treated with LMB for 22 hours and additionally with the different secretase inhibitors as described in C. Cells were stained as in B and single confocal sections were acquired in the mid-nuclear plane as determined by the Map2ab staining. The upper row shows an overlay of all three stainings whereas the lower row is a twofold zoom of the APP staining. Treatment with either LMB alone or in combination with TAPI1 results in a relatively high frequency of AICD nuclear spots. By contrast, treatment of neurons with DAPT or either BACE1 inhibitor A or B reduces the frequency of nuclear AICD spots. Scale bar: 7 μm. (E) Quantification of the number of AICD spots per nucleus reveals a significant reduction in the number of nuclear AICD spots following treatment with either DAPT or BACE1 inhibitors. <0.001 versus LMB alone, MWU. Treatment of cells with TAPI 1 does not significantly alter the number of nuclear AICD spots, when compared to LMB treatment alone. Number of imaged neurons: control, =19; LMB, =35; DAPT+LMB, =34; TAPI 1+LMB, =42; BACE1 inhibitor A+LMB, =38; BACE1 inhibitor B+LMB, =22, error bars represent s.e.m.

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The physiological outcomes of α- or β-secretase-initiated processing of APP are distinct. We have now shown that this dichotomy holds up for AICD nuclear signaling. The β-secretase-mediated cleavage of APP in endosomal compartments is mainly responsible for generating AICD species that are capable of translocating to the nucleus, and active transport of endosomes to the perinuclear region is a prerequisite for nuclear signaling.

Banks that were found to have problems "have already taken -- or are in the process of" taking corrective actions such as closing accounts, refunding customers and fixing credit bureau information, the OCC said.

"It's certainly troubling that this appears to be more widespread than one bank," said John Sedunov, a finance professor at Villanova.

A PwC consultant briefed on the findings told the American Banker that the review led the OCC to issue 252 alerts to banks about matters that required attention as well as five industry-wide problems to be fixed.

The OCC declined to comment beyond its statement. PwC declined to comment.

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Wells Fargo ( ) sent shockwaves through the industry in September 2016 when it admitted to firing 5,300 workers for opening millions of fake accounts.

It quickly emerged that wildly unrealistic sales goals and unrelenting pressure from management led employees to open accounts for customers who didn't want or need them. Some Wells Fargo bankers impersonated their customers and used false email addresses like noname@wellsfargo.com, according to a 2015 lawsuit filed by the city of Los Angeles.

But even then there were signs Wells Fargo wasn't alone. Nearly a dozen current and former employees at large and regional banks such as Bank of America ( ) , Citizens Bank ( ) , PNC ( ) and SunTrust ( ) told CNNMoney in September 2016 that a sales obsession pervaded their banks too. They described immense pressure to get customers to open multiple accounts.

"Wells Fargo isn't the only one. This is an industry-wide problem," one former banker at a regional lender told CNNMoney.

The OCC said it completed its review in late 2017 and letters to participating banks were sent earlier this year. The agency supervises all national banks, including major lenders such as JPMorgan Chase ( ) , Citigroup ( ) , Bank of America, US Bank ( ) and PNC. The OCC also regulates federal branches of foreign banks.

A PNC spokesperson said the bank regularly trains employees to have "thoughtful discussions" with customers about financial products. "We only act on what a customer wants to pursue," PNC said.

Citigroup ( ) , US Bank and Bank of America declined to comment.

Related: Wells Fargo altered documents about business clients

The OCC does not plan to publish a report on its review of bank sales tactics, meaning the scale of the troubles remain a mystery.