{
  "abstract": "A fundamental but unanswered biological question is how much energy life forms expend per unit mass and time to remain alive (Makarieva et al., 2008). The West-Brown-Enquist (WBE) theory explains Kleiber's 3/4-power law (Kleiber, 1932) through complex vascular networks (West et al., 1997; Banavar et al., 1999; West et al., 1999; Dodds et al., 2001; Gillooly et al., 2001; West et al., 2002; Banavar et al., 2002; Kozowski & Konarzewski, 2004; Brown et al., 2005; White et al., 2007; Savage et al., 2008; Apol et al., 2008), but assumes \"size-invariant terminal units\" and predicts neither absolute rates, small-organism linearity, nor group-specific levels. Yet capillary dimensions vary with body mass (Dawson, 2003) and obey Murray's optimization principle (Murray, 1926). Furthermore, 3/4-power scaling occurs in planarians with a simple tubular metabolic system (Thommen et al., 2019). Here we extend Krogh's capillary--tissue unit (Krogh, 1919) to an optimized mesoscale oxygen-transport system, the Metabolic Holon (MH). Krogh's geometry, Murray's minimum-work principle, and optimal oxygen utilization determine its geometry and operating state. Kleiber's asymptote thus emerges from an optimized Krogh unit, without requiring complex vascular-network architecture; repetition and within-group body-mass-invariant topology determine organismal metabolism. Without fitting metabolic-rate data, MH predicts absolute rates across 18 orders of magnitude (R^2 = 97.7 %), including linear scaling and group-specific Kleiber asymptotes. Independent dimensional analysis recovers these asymptotes. The theory also predicts absolute mammalian heart rates without calibration; absolute lifespan additionally requires an empirical lifetime-heartbeat constraint. By representing evolutionary selection and adaptation through mesoscale optimization, the theory explains why \"mass-specific metabolic rates are strikingly similar across life's major domains\" (Makarieva et al., 2008).",
  "abstract_shown": true,
  "affiliations": [
    "Fluidsystemtechnik, TU Darmstadt"
  ],
  "article_type": "New results",
  "arxiv": null,
  "author_total": 2,
  "authors": [
    {
      "affs": [
        0
      ],
      "name": "P. F. Pelz"
    },
    {
      "name": "T. C. Meck"
    }
  ],
  "bibtex": "https://research.draconitic.com/paper/a7p88qkkm5k74.bib",
  "cited_by": 0,
  "doi": "10.64898/2026.09.03.749078",
  "fulltext_url": "https://www.biorxiv.org/content/10.64898/2026.09.03.749078v1.full",
  "id": "a7p88qkkm5k74",
  "issns": [],
  "landing_url": "https://www.biorxiv.org/content/10.64898/2026.09.03.749078v1",
  "language": null,
  "licence": "CC-BY",
  "licence_label": "CC BY",
  "licence_source": "article",
  "licence_url": "https://creativecommons.org/licenses/by/4.0/",
  "metadata_licence": null,
  "native_subjects": [
    [
      "biorxiv",
      "physiology"
    ],
    [
      "openalex",
      "2303|Physiological and biochemical adaptations"
    ]
  ],
  "openalex": "W7211990744",
  "pdf_url": "https://www.biorxiv.org/content/10.64898/2026.09.03.749078v1.full.pdf",
  "pmcid": null,
  "pmid": null,
  "published": "2026-09-03",
  "publisher": null,
  "ris": "https://research.draconitic.com/paper/a7p88qkkm5k74.ris",
  "sources": [
    {
      "first_seen": "2026-10-30T12:20:38Z",
      "native_id": "10.64898/2026.09.03.749078",
      "source": "biorxiv",
      "url": "https://www.biorxiv.org/content/10.64898/2026.09.03.749078v4"
    }
  ],
  "subjects": [
    {
      "field": "medicine",
      "origin": "biorxiv:physiology",
      "subfield": "physiology"
    }
  ],
  "title": "Local optimization of oxygen transport gives rise to Kleiber's law",
  "title_text": "Local optimization of oxygen transport gives rise to Kleiber's law",
  "type": "preprint",
  "updated_at": "2026-10-30T12:29:19Z",
  "url": "https://research.draconitic.com/paper/a7p88qkkm5k74",
  "venue": "bioRxiv",
  "versions": []
}