---
_id: '12232'
abstract:
- lang: eng
  text: We investigate the "macronutrient-access hypothesis", which states that the
    balance between stoichiometric macronutrient demand and accessible macronutrients
    controls nutrient assimilation by aquatic heterotrophs. Within this hypothesis,
    we consider bioavailable dissolved organic carbon (bDOC), reactive nitrogen (N)
    and reactive phosphorus (P) to be the macronutrients accessible to heterotrophic
    assimilation. Here, reactive N and P are the sums of dissolved inorganic N (nitrate-N,
    nitrite-N, ammonium-N), soluble-reactive P (SRP), and bioavailable dissolved organic
    N (bDON) and P (bDOP). Previous data from various freshwaters suggests this hypothesis,
    yet clear experimental support is missing. We assessed this hypothesis in a proof-of-concept
    experiment for waters from four small agricultural streams. We used seven different
    bDOC:reactive N and bDOC:reactive P ratios, induced by seven levels of alder leaf
    leachate addition. With these treatments and a stream-water specific bacterial
    inoculum, we conducted a 3-day experiment with three independent replicates per
    combination of stream water, treatment, and sampling occasion. Here, we extracted
    dissolved organic matter (DOM) fluorophores by measuring excitation-emission matrices
    with subsequent parallel factor decomposition (EEM-PARAFAC). We assessed the true
    bioavailability of DOC, DON, and the DOM fluorophores as the concentration difference
    between the beginning and end of each experiment. Subsequently, we calculated
    the bDOC and bDON concentrations based on the bioavailable EEM-PARAFAC fluorophores,
    and compared the calculated bDOC and bDON concentrations to their true bioavailability.
    Due to very low DOP concentrations, the DOP determination uncertainty was high,
    and we assumed DOP to be a negligible part of the reactive P. For bDOC and bDON,
    the true bioavailability measurements agreed with the same fractions calculated
    indirectly from bioavailable EEM-PARAFAC fluorophores (bDOC r2 = 0.96, p < 0.001;
    bDON r2 = 0.77, p < 0.001). Hence we could predict bDOC and bDON concentrations
    based on the EEM-PARAFAC fluorophores. The ratios of bDOC:reactive N (sum of bDON
    and DIN) and bDOC:reactive P (equal to SRP) exerted a strong, predictable stoichiometric
    control on reactive N and P uptake (R2 = 0.80 and 0.83). To define zones of C:N:P
    (co-)limitation of heterotrophic assimilation, we used a novel ternary-plot approach
    combining our data with literature data on C:N:P ranges of bacterial biomass.
    Here, we found a zone of maximum reactive N uptake (C:N:P approx. > 114: < 9:1),
    reactive P uptake (C:N:P approx. > 170:21: < 1) and reactive N and P co-limitation
    of nutrient uptake (C:N:P approx. > 204:14:1). The “macronutrient-access hypothesis”
    links ecological stoichiometry and biogeochemistry, and may be of importance for
    nutrient uptake in many freshwater ecosystems. However, this experiment is only
    a starting point and this hypothesis needs to be corroborated by further experiments
    for more sites, by in-situ studies, and with different DOC sources.
author:
- first_name: Daniel
  full_name: Graeber, Daniel
  last_name: Graeber
- first_name: Youngdoung
  full_name: Tenzin, Youngdoung
  last_name: Tenzin
- first_name: Marc
  full_name: Stutter, Marc
  last_name: Stutter
- first_name: Gabriele
  full_name: Weigelhofer, Gabriele
  last_name: Weigelhofer
- first_name: Tom
  full_name: Shatwell, Tom
  id: '86424'
  last_name: Shatwell
  orcid: 0000-0002-4520-7916
- first_name: Wolf
  full_name: von Tümpling, Wolf
  last_name: von Tümpling
- first_name: Jörg
  full_name: Tittel, Jörg
  last_name: Tittel
- first_name: Alexander
  full_name: Wachholz, Alexander
  last_name: Wachholz
- first_name: Dietrich
  full_name: Borchardt, Dietrich
  last_name: Borchardt
citation:
  ama: 'Graeber D, Tenzin Y, Stutter M, et al. Bioavailable DOC: reactive nutrient
    ratios control heterotrophic nutrient assimilation—An experimental proof of the
    macronutrient-access hypothesis. <i>Biogeochemistry : an international journal
    </i>. 2021;155(1):1-20. doi:<a href="https://doi.org/10.1007/s10533-021-00809-4">10.1007/s10533-021-00809-4</a>'
  apa: 'Graeber, D., Tenzin, Y., Stutter, M., Weigelhofer, G., Shatwell, T., von Tümpling,
    W., Tittel, J., Wachholz, A., &#38; Borchardt, D. (2021). Bioavailable DOC: reactive
    nutrient ratios control heterotrophic nutrient assimilation—An experimental proof
    of the macronutrient-access hypothesis. <i>Biogeochemistry : An International
    Journal </i>, <i>155</i>(1), 1–20. <a href="https://doi.org/10.1007/s10533-021-00809-4">https://doi.org/10.1007/s10533-021-00809-4</a>'
  bjps: '<b>Graeber D <i>et al.</i></b> (2021) Bioavailable DOC: Reactive Nutrient
    Ratios Control Heterotrophic Nutrient Assimilation—An Experimental Proof of the
    Macronutrient-Access Hypothesis. <i>Biogeochemistry : an international journal
    </i> <b>155</b>, 1–20.'
  chicago: 'Graeber, Daniel, Youngdoung Tenzin, Marc Stutter, Gabriele Weigelhofer,
    Tom Shatwell, Wolf von Tümpling, Jörg Tittel, Alexander Wachholz, and Dietrich
    Borchardt. “Bioavailable DOC: Reactive Nutrient Ratios Control Heterotrophic Nutrient
    Assimilation—An Experimental Proof of the Macronutrient-Access Hypothesis.” <i>Biogeochemistry :
    An International Journal </i> 155, no. 1 (2021): 1–20. <a href="https://doi.org/10.1007/s10533-021-00809-4">https://doi.org/10.1007/s10533-021-00809-4</a>.'
  chicago-de: 'Graeber, Daniel, Youngdoung Tenzin, Marc Stutter, Gabriele Weigelhofer,
    Tom Shatwell, Wolf von Tümpling, Jörg Tittel, Alexander Wachholz und Dietrich
    Borchardt. 2021. Bioavailable DOC: reactive nutrient ratios control heterotrophic
    nutrient assimilation—An experimental proof of the macronutrient-access hypothesis.
    <i>Biogeochemistry : an international journal </i> 155, Nr. 1: 1–20. doi:<a href="https://doi.org/10.1007/s10533-021-00809-4">10.1007/s10533-021-00809-4</a>,
    .'
  din1505-2-1: '<span style="font-variant:small-caps;"><span style="font-variant:small-caps;">Graeber,
    Daniel</span> ; <span style="font-variant:small-caps;">Tenzin, Youngdoung</span>
    ; <span style="font-variant:small-caps;">Stutter, Marc</span> ; <span style="font-variant:small-caps;">Weigelhofer,
    Gabriele</span> ; <span style="font-variant:small-caps;">Shatwell, Tom</span>
    ; <span style="font-variant:small-caps;">von Tümpling, Wolf</span> ; <span style="font-variant:small-caps;">Tittel,
    Jörg</span> ; <span style="font-variant:small-caps;">Wachholz, Alexander</span>
    ; u. a.</span>: Bioavailable DOC: reactive nutrient ratios control heterotrophic
    nutrient assimilation—An experimental proof of the macronutrient-access hypothesis.
    In: <i>Biogeochemistry : an international journal </i> Bd. 155. Cham, Springer  (2021),
    Nr. 1, S. 1–20'
  havard: 'D. Graeber, Y. Tenzin, M. Stutter, G. Weigelhofer, T. Shatwell, W. von
    Tümpling, J. Tittel, A. Wachholz, D. Borchardt, Bioavailable DOC: reactive nutrient
    ratios control heterotrophic nutrient assimilation—An experimental proof of the
    macronutrient-access hypothesis, Biogeochemistry : An International Journal .
    155 (2021) 1–20.'
  ieee: 'D. Graeber <i>et al.</i>, “Bioavailable DOC: reactive nutrient ratios control
    heterotrophic nutrient assimilation—An experimental proof of the macronutrient-access
    hypothesis,” <i>Biogeochemistry : an international journal </i>, vol. 155, no.
    1, pp. 1–20, 2021, doi: <a href="https://doi.org/10.1007/s10533-021-00809-4">10.1007/s10533-021-00809-4</a>.'
  mla: 'Graeber, Daniel, et al. “Bioavailable DOC: Reactive Nutrient Ratios Control
    Heterotrophic Nutrient Assimilation—An Experimental Proof of the Macronutrient-Access
    Hypothesis.” <i>Biogeochemistry : An International Journal </i>, vol. 155, no.
    1, 2021, pp. 1–20, <a href="https://doi.org/10.1007/s10533-021-00809-4">https://doi.org/10.1007/s10533-021-00809-4</a>.'
  short: 'D. Graeber, Y. Tenzin, M. Stutter, G. Weigelhofer, T. Shatwell, W. von Tümpling,
    J. Tittel, A. Wachholz, D. Borchardt, Biogeochemistry : An International Journal  155
    (2021) 1–20.'
  ufg: '<b>Graeber, Daniel u. a.</b>: Bioavailable DOC: reactive nutrient ratios control
    heterotrophic nutrient assimilation—An experimental proof of the macronutrient-access
    hypothesis, in: <i>Biogeochemistry : an international journal </i> 155 (2021),
    H. 1,  S. 1–20.'
  van: 'Graeber D, Tenzin Y, Stutter M, Weigelhofer G, Shatwell T, von Tümpling W,
    et al. Bioavailable DOC: reactive nutrient ratios control heterotrophic nutrient
    assimilation—An experimental proof of the macronutrient-access hypothesis. Biogeochemistry :
    an international journal . 2021;155(1):1–20.'
date_created: 2024-12-08T20:21:39Z
date_updated: 2024-12-09T11:21:31Z
department:
- _id: DEP8022
doi: 10.1007/s10533-021-00809-4
extern: '1'
intvolume: '       155'
issue: '1'
language:
- iso: eng
main_file_link:
- open_access: '1'
  url: https://doi.org/10.1007/s10533-021-00809-4
oa: '1'
page: 1-20
place: Cham
publication: 'Biogeochemistry : an international journal '
publication_identifier:
  eissn:
  - 1573-515X
  issn:
  - 0168-2563
publication_status: published
publisher: 'Springer '
quality_controlled: '1'
status: public
title: 'Bioavailable DOC: reactive nutrient ratios control heterotrophic nutrient
  assimilation—An experimental proof of the macronutrient-access hypothesis'
type: scientific_journal_article
user_id: '83781'
volume: 155
year: '2021'
...
