Silicon, endophytes and secondary metabolites as grass defenses against mammalian herbivores

Grasses have been considered to primarily employ tolerance in lieu of defense in mitigating damage caused by herbivory. Yet a number of mechanisms have been identified in grasses, which may deter feeding by grazers. These include enhanced silicon uptake, hosting of toxin-producing endophytic fungi a...

Full description

Bibliographic Details
Main Authors: Otso eHuitu, Kristian eForbes, Marjo eHelander, Riitta eJulkunen-Tiitto, Xavier eLambin, Kari eSaikkonen, Peter eStuart, Sini eSulkama, Sue Elaine Hartley
Format: Article
Language:English
Published: Frontiers Media S.A. 2014-09-01
Series:Frontiers in Plant Science
Subjects:
Online Access:http://journal.frontiersin.org/Journal/10.3389/fpls.2014.00478/full
_version_ 1819327168793018368
author Otso eHuitu
Kristian eForbes
Marjo eHelander
Riitta eJulkunen-Tiitto
Xavier eLambin
Kari eSaikkonen
Peter eStuart
Sini eSulkama
Sue Elaine Hartley
author_facet Otso eHuitu
Kristian eForbes
Marjo eHelander
Riitta eJulkunen-Tiitto
Xavier eLambin
Kari eSaikkonen
Peter eStuart
Sini eSulkama
Sue Elaine Hartley
author_sort Otso eHuitu
collection DOAJ
description Grasses have been considered to primarily employ tolerance in lieu of defense in mitigating damage caused by herbivory. Yet a number of mechanisms have been identified in grasses, which may deter feeding by grazers. These include enhanced silicon uptake, hosting of toxin-producing endophytic fungi and induction of secondary metabolites. While these mechanisms have been individually studied, their synergistic responses to grazing, as well as their effects on grazers, are poorly known. A field experiment was carried out in 5 × 5 m outdoor enclosures to quantify phytochemical changes of either endophyte-infected (E+) or endophyte-free (E-) meadow fescue (Schedonorus pratensis) in response to medium intensity (corresponding with densities of ca. 1200 voles / ha for 5 weeks during 3 months) or heavy intensity (ca. 1200 voles / ha for 8 weeks during 3 months) grazing by a mammalian herbivore, the field vole (Microtus agrestis). A laboratory experiment was then conducted to evaluate the effects of endophyte infection status and grazing history of the grass diet on vole performance. As predicted, grazing increased foliar silicon content, by up to 13 %. Grazing also increased foliar levels of phosphorous and several phenolic compounds, most notably those of the flavonols isorhamnetin-diglycoside and rhamnetin derivative. Silicon concentrations were consistently circa 16 % higher in E+ grasses than in E-grasses, at all levels of grazing. Similarly, concentrations of chlorogenic acid derivative were found to be consistently higher in E+ than in E- grasses. Female voles maintained on heavily grazed grasses suffered higher mortality rates in the laboratory than female voles fed ungrazed grass, regardless of endophyte infection status. Our results conclusively demonstrate that, in addition to tolerance, grasses employ multi-tiered, effective defenses against mammalian grazers.
first_indexed 2024-12-24T13:06:33Z
format Article
id doaj.art-4d1e9e80d49d4a0790aedf60cbed9c21
institution Directory Open Access Journal
issn 1664-462X
language English
last_indexed 2024-12-24T13:06:33Z
publishDate 2014-09-01
publisher Frontiers Media S.A.
record_format Article
series Frontiers in Plant Science
spelling doaj.art-4d1e9e80d49d4a0790aedf60cbed9c212022-12-21T16:53:59ZengFrontiers Media S.A.Frontiers in Plant Science1664-462X2014-09-01510.3389/fpls.2014.00478100750Silicon, endophytes and secondary metabolites as grass defenses against mammalian herbivoresOtso eHuitu0Kristian eForbes1Marjo eHelander2Riitta eJulkunen-Tiitto3Xavier eLambin4Kari eSaikkonen5Peter eStuart6Sini eSulkama7Sue Elaine Hartley8Finnish Forest Research InstituteFinnish Forest Research InstituteUniversity of TurkuUniversity of Eastern FinlandUniversity of AberdeenAgrifood Research FinlandFinnish Forest Research InstituteFinnish Forest Research InstituteUniversity of YorkGrasses have been considered to primarily employ tolerance in lieu of defense in mitigating damage caused by herbivory. Yet a number of mechanisms have been identified in grasses, which may deter feeding by grazers. These include enhanced silicon uptake, hosting of toxin-producing endophytic fungi and induction of secondary metabolites. While these mechanisms have been individually studied, their synergistic responses to grazing, as well as their effects on grazers, are poorly known. A field experiment was carried out in 5 × 5 m outdoor enclosures to quantify phytochemical changes of either endophyte-infected (E+) or endophyte-free (E-) meadow fescue (Schedonorus pratensis) in response to medium intensity (corresponding with densities of ca. 1200 voles / ha for 5 weeks during 3 months) or heavy intensity (ca. 1200 voles / ha for 8 weeks during 3 months) grazing by a mammalian herbivore, the field vole (Microtus agrestis). A laboratory experiment was then conducted to evaluate the effects of endophyte infection status and grazing history of the grass diet on vole performance. As predicted, grazing increased foliar silicon content, by up to 13 %. Grazing also increased foliar levels of phosphorous and several phenolic compounds, most notably those of the flavonols isorhamnetin-diglycoside and rhamnetin derivative. Silicon concentrations were consistently circa 16 % higher in E+ grasses than in E-grasses, at all levels of grazing. Similarly, concentrations of chlorogenic acid derivative were found to be consistently higher in E+ than in E- grasses. Female voles maintained on heavily grazed grasses suffered higher mortality rates in the laboratory than female voles fed ungrazed grass, regardless of endophyte infection status. Our results conclusively demonstrate that, in addition to tolerance, grasses employ multi-tiered, effective defenses against mammalian grazers.http://journal.frontiersin.org/Journal/10.3389/fpls.2014.00478/fullEndophytesSilicongrassesDefensegrazingphenolics
spellingShingle Otso eHuitu
Kristian eForbes
Marjo eHelander
Riitta eJulkunen-Tiitto
Xavier eLambin
Kari eSaikkonen
Peter eStuart
Sini eSulkama
Sue Elaine Hartley
Silicon, endophytes and secondary metabolites as grass defenses against mammalian herbivores
Frontiers in Plant Science
Endophytes
Silicon
grasses
Defense
grazing
phenolics
title Silicon, endophytes and secondary metabolites as grass defenses against mammalian herbivores
title_full Silicon, endophytes and secondary metabolites as grass defenses against mammalian herbivores
title_fullStr Silicon, endophytes and secondary metabolites as grass defenses against mammalian herbivores
title_full_unstemmed Silicon, endophytes and secondary metabolites as grass defenses against mammalian herbivores
title_short Silicon, endophytes and secondary metabolites as grass defenses against mammalian herbivores
title_sort silicon endophytes and secondary metabolites as grass defenses against mammalian herbivores
topic Endophytes
Silicon
grasses
Defense
grazing
phenolics
url http://journal.frontiersin.org/Journal/10.3389/fpls.2014.00478/full
work_keys_str_mv AT otsoehuitu siliconendophytesandsecondarymetabolitesasgrassdefensesagainstmammalianherbivores
AT kristianeforbes siliconendophytesandsecondarymetabolitesasgrassdefensesagainstmammalianherbivores
AT marjoehelander siliconendophytesandsecondarymetabolitesasgrassdefensesagainstmammalianherbivores
AT riittaejulkunentiitto siliconendophytesandsecondarymetabolitesasgrassdefensesagainstmammalianherbivores
AT xavierelambin siliconendophytesandsecondarymetabolitesasgrassdefensesagainstmammalianherbivores
AT kariesaikkonen siliconendophytesandsecondarymetabolitesasgrassdefensesagainstmammalianherbivores
AT peterestuart siliconendophytesandsecondarymetabolitesasgrassdefensesagainstmammalianherbivores
AT siniesulkama siliconendophytesandsecondarymetabolitesasgrassdefensesagainstmammalianherbivores
AT sueelainehartley siliconendophytesandsecondarymetabolitesasgrassdefensesagainstmammalianherbivores