# Forage quality shapes physiological and gut microbial responses in moose (Alces alces) of Isle Royale National Park

**Authors:** Sebastian Menke, Gloria Fackelmann, Leah M. Vucetich, John A. Vucetich, Jennifer S. Forbey, Simone Sommer

PMC · DOI: 10.1038/s41598-026-35555-w · 2026-01-27

## TL;DR

Moose on Isle Royale eat balsam fir in winter, which affects their health and gut microbes, showing how diet shapes both.

## Contribution

This study reveals how moose adapt to a chemically defended diet through physiological and microbial changes.

## Key findings

- Balsam fir consumption increases detoxification markers and reduces nutritional condition in moose.
- Higher fir intake correlates with lower gut microbial diversity and shifts in bacterial genera like Roseburia.
- Metagenomic analysis shows pathways for degrading plant secondary compounds, though not all are significantly linked to fir consumption.

## Abstract

Plant secondary compounds (PSCs) impose physiological and nutritional constraints on herbivores, yet many species continue to rely on PSC-rich forage during critical periods of the year. Moose (Alces alces) on Isle Royale National Park depend heavily on balsam fir (Abies balsamea) during winter, exposing them to a chemically defended and nutritionally limited diet. To investigate how this foraging strategy shapes both physiological responses and gut microbial communities, we integrated fecal diet composition, urinary markers of detoxification and nutritional status, 16S rRNA gene sequencing, and shotgun metagenomic profiling from free ranging moose of two separated populations from the western and eastern region of the island. Balsam fir consumption varied strongly by region and was positively associated with glucuronic acid to creatinine (GA:C) and urea nitrogen to creatinine (UN:C) ratios, indicating increased detoxification activity and reduced nutritional condition. Microbial alpha diversity declined with higher fir intake in nutritionally limited individuals, while beta diversity differed by region, balsam fir consumption, and UN:C. Several bacterial genera responded to PSC exposure, including increases in the butyrate-producing genus Roseburia and shifts in network prominence of Phascolarctobacterium. Metagenomic data revealed pathways involved in the degradation of aromatic and terpenoid PSCs, although pathway abundances did not differ significantly with balsam fir consumption after multiple testing correction. These results show that winter foraging on balsam fir produces coordinated dietary, physiological, and microbial patterns, with both host and gut microbial detoxification capacities interacting to accommodate the chemical and nutritional challenges of a PSC-rich winter diet.

The online version contains supplementary material available at 10.1038/s41598-026-35555-w.

## Linked entities

- **Chemicals:** glucuronic acid (PubChem CID 65041), urea nitrogen (PubChem CID 31295), creatinine (PubChem CID 588)
- **Species:** Alces alces (taxon 9852), Abies balsamea (taxon 90345), Mus musculus (taxon 10090)

## Full-text entities

- **Diseases:** bacterial (MESH:D001424), carcinogenic (MESH:D011230), PSC (MESH:D010939), neurotoxicity (MESH:D020258), death (MESH:D003643), starvation (MESH:D013217)
- **Chemicals:** thymol (MESH:D013943), Polycyclic aromatic hydrocarbon (MESH:D011084), Cofactors (MESH:C013123), Amino Acid (MESH:D000596), Dioxin (MESH:D004147), Polyketides (MESH:D061065), chlorobenzene (MESH:C031294), Butyrate (MESH:D002087), Carbazole (MESH:C041514), camphene (MESH:C019286), Aminobenzoate (MESH:D062365), nitrogen (MESH:D009584), Benzoate (MESH:D001565), NaOH (MESH:D012972), ansamycins (MESH:D047029), Tetracycline (MESH:D013752), Carotenoid (MESH:D002338), Xylene (MESH:D014992), monoterpenes (MESH:D039821), UN (MESH:C530477), SCFA (MESH:D005232), vancomycin (MESH:D014640), citrate (MESH:D019343), PSC (-), creatinine (MESH:D003404), piperitone (MESH:C496656), Styrene (MESH:D020058), tannin (MESH:D013634), urea (MESH:D014508), Terpenoid (MESH:D013729), GA (MESH:D020723), 1,2-Dichloroethane (MESH:C024565), CoA (MESH:D003065), Carbohydrate (MESH:D002241), CF (MESH:D002142), C (MESH:D002244), beta bisabolene (MESH:C531191), Delta3 carene (MESH:C030218), Limonene (MESH:D000077222), alkaloids (MESH:D000470), Fluorene (MESH:C041509), Zeatin (MESH:D015026), H2O (MESH:D014867), longifolene (MESH:C035607), ornithine (MESH:D009952), bornyl acetate (MESH:C071528), acetyl-CoA (MESH:D000105), alpha terpineol (MESH:C016775), beta-pinene (MESH:C010789)
- **Species:** Bonasa umbellus (ruffed grouse, species) [taxon 9000], Alces americanus (American moose, species) [taxon 999462], Juniperus communis (common juniper, species) [taxon 58039], Faecalibacterium (genus) [taxon 216851], Prevotella (genus) [taxon 838], Phascolarctobacterium (genus) [taxon 33024], Nocardioides (genus) [taxon 1839], Roseburia (genus) [taxon 841], Odocoileus hemionus (mule deer, species) [taxon 9872], Odocoileus virginianus (white-tailed deer, species) [taxon 9874], Mycoplasma (genus) [taxon 2093], Juniperus monosperma (cherrystone juniper, species) [taxon 487038], Psittacidae (parrot, family) [taxon 9224], Clostridium (genus) [taxon 1485], Abies balsamea (balsam fir, species) [taxon 90345], Neotoma albigula (white-throated woodrat, species) [taxon 42408], Neotoma stephensi (species) [taxon 164611], Actinomycetospora (genus) [taxon 402649], Bos taurus (bovine, species) [taxon 9913], gut metagenome (species) [taxon 749906], Bacillota (clostridial firmicutes, phylum) [taxon 1239], Methyloversatilis (genus) [taxon 378210], Alces alces (elk, species) [taxon 9852], Bacteroides (genus) [taxon 816], Ruminococcus (genus) [taxon 1263], Homo sapiens (human, species) [taxon 9606], Coprococcus (genus) [taxon 33042]

## Figures

11 figures with captions in the complete paper: https://tomesphere.com/paper/PMC12852835/full.md

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Source: https://tomesphere.com/paper/PMC12852835