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We are perceptually most sensitive to slow modulations which are most common. However, previous studies of envelope coding in the brainstem found an under-representation of these slow, low-frequency, modulations. Specifically, the synchronization of spike times to the envelope was enhanced in some neuron types, forming channels specialized for envelope processing but tuned to a restricted range of fast, high-frequency, envelopes (200\u2013500 Hz). Here, we show using a historical dataset from cats that previous analyses, which made strong assumptions about the neural code, underestimated the encoding of low-frequency envelopes. While some neurons encode envelope better than others, most encode a wide range of envelope frequencies, and represent slower envelope fluctuations most accurately in their precise patterns of spike times. Identification of envelope frequency from spike-timing was linked to reliability, and to the way that dynamics of spiking interacted with the time-varying envelope. In some of the best-performing neurons, temporally complex \u201cmode-locked\u201d spike patterns served to enhance envelope coding. A second long-standing contradiction was that neural envelope coding is degraded at high sound levels, whilst the perception of envelope is robust at a wide range of sound levels. We find that spike-time encoding of envelope shape becomes level-robust for small populations of neurons. These findings argue against feature-specific coding of envelopes in the brainstem, and for a distributed population spike-time code for which synchrony to the envelope is an incomplete description. This code is accurate for slow fluctuations and robust across sound level. Thus, precise spike-timing information in the brainstem is after-all aligned with the needs of communication and the perception of environmental sounds.<\/jats:p>","DOI":"10.1371\/journal.pbio.3003213","type":"journal-article","created":{"date-parts":[[2025,6,16]],"date-time":"2025-06-16T17:38:48Z","timestamp":1750095528000},"page":"e3003213","update-policy":"https:\/\/doi.org\/10.1371\/journal.pbio.corrections_policy","source":"Crossref","is-referenced-by-count":1,"title":["Precise spike-timing information in the brainstem is well aligned with the needs of communication and the perception of environmental sounds"],"prefix":"10.1371","volume":"23","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1597-3070","authenticated-orcid":true,"given":"Chris","family":"Scholes","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Stephen","family":"Coombes","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Alan R.","family":"Palmer","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"William S.","family":"Rhode","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Rob","family":"Mill","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2573-7418","authenticated-orcid":true,"given":"Christian J.","family":"Sumner","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"340","published-online":{"date-parts":[[2025,6,16]]},"reference":[{"issue":"5234","key":"pbio.3003213.ref001","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1126\/science.270.5234.303","article-title":"Speech recognition with primarily temporal cues","volume":"270","author":"RV Shannon","year":"1995","journal-title":"Science"},{"issue":"4","key":"pbio.3003213.ref002","doi-asserted-by":"crossref","first-page":"2403","DOI":"10.1121\/1.419603","article-title":"Speech intelligibility as a function of the number of channels of stimulation for signal processors using sine-wave and noise-band outputs","volume":"102","author":"MF Dorman","year":"1997","journal-title":"J Acoust Soc Am"},{"key":"pbio.3003213.ref003","doi-asserted-by":"crossref","first-page":"3394","DOI":"10.1121\/1.1624067","article-title":"Modulation spectra of natural sounds and ethological theories of auditory processing","volume":"114","author":"NC Singh","year":"2003","journal-title":"J Acoust Soc Am"},{"issue":"1","key":"pbio.3003213.ref004","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1121\/1.402757","article-title":"Responses to amplitude-modulated tones in the auditory nerve of the cat","volume":"91","author":"PX Joris","year":"1992","journal-title":"J Acoust Soc Am"},{"key":"pbio.3003213.ref005","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/0378-5955(90)90074-Y","article-title":"Encoding of amplitude modulation in the gerbil cochlear nucleus: I. 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