FUN（newX [，i]，…）中的错误：参数“ x”丢失，没有默认值

我正在尝试将原始计数转换为R中的fpkm。

expMatrix = read.csv("counts.csv", header = TRUE, row.names = 1, check.names = FALSE)
countToTpm <- function(counts, effLen)
{
  rate <- log(counts) - log(effLen)
  denom <- log(sum(exp(rate)))
  exp(rate - denom + log(1e6))
}

countToFpkm <- function(counts, effLen)
{
  N <- sum(counts)
  exp( log(counts) + log(1e9) - log(effLen) - log(N) )
}

fpkmToTpm <- function(fpkm)
{
  exp(log(fpkm) - log(sum(fpkm)) + log(1e6))
}

countToEffCounts <- function(counts, len, effLen)
{
  counts * (len / effLen)
}

tpms <- apply(expMatrix,2,countToFpkm)

FUN（newX [，i]，...）中的错误：缺少参数“ x”，没有默认值

有谁知道如何解决这个问题？