original research article
submitted: 09.09.2015, reviewed: 13.10.2015, accepted after revision: 30.10.2015.
© 2016 a. Klisz et al.
e-ISSN 2082-8926
The vulnerability of silver fir populations to damage from late frosts
Marcin Klisz1*, Szymon Jastrzębowski1, Joanna Ukalska2, Paweł Przybylski1, Jan Matras1, Marcin Mionskowski3
1 orest esearc nstit te, e art ent o il ic lt re and enetics o orest rees, l. Braci Leśne 3, ocin tar , 0 –0 0 asz n, Poland; 2 arsaw ni ersit Li e ciences – , e art ent o cono etrics and tatistics, i ision o Bio etrics, l. owo rs nows a 1 , 02–776 arszawa, Poland 3Forest research institute, Department of Forest resources Management,
l. Braci Leśne 3, ocin tar , 0 –0 0 asz n, Poland el. 4 22 71 0 3 2, a 4 22 72003 7, e ail: . lisz i les.waw. l
Abstract. e ai o t e st d was to deter ine t e lnera ilit o selected sil er r o lations to da a e ro late rost in t e cli atic conditions o so t eastern Poland. o deter ine t e lnera ilit o a ical and lateral s oots to da a e ca sed late rosts, we o ser ed o r test lots in 200 and 2014, eac containin ro enies o selected seed stands. r statistical anal ses were ased on a odel incor oratin t e ollowin aria les: site, ear, t e o rost da a e, o lation as well as t e ossi le interaction etween t ese aria les. i ni cant di erences etween t e o lations were o nd in ter s o t eir sensiti it to da a e ro low te erat re occ rrin d rin t e rowt eriod. rt er ore, we indirectl de onstrated di erences in t e se erit o late rost on t e e eri ental lots, as well as t e intensit and aria ilit o late rost s oot da a e. Based on t ese res lts, we di ided t e st died o lations into two ro s o low ( , K A1 and A ) and i (L 2 and BAL2) sensiti it to late frost damage.
Keywords: adaptation, environmental stress, frost damage, testing program, Abies alba
1. Introduction
n c oosin orest trees, testin t e ro en o selected o lations and enot es in eld e eri ents la s a e role in understanding their adaptive potential in the context of predicted and observed climate changes (ledig, Kitzmiller 1 2 anewin el et al. 2012). estin o lations o orest trees el s to deter ine t eir ada ti it , direction and sco e o c an e, as well as to e al ate t e e i ilit o t e ro e- nance in a aria le en iron ent (Ait en, itloc 2013).
e earliest atte ts to de ne t e enetic aria ilit o ada- ptive traits of forest trees based on experiments of scots pine
ro enances were ade P ili e Pierre Andre de Vil o- rin in t e rst al o t e nineteent cent r (see Lan let 1 71). owe er, t e rst eld e eri ents sin a la o t for provenance trials with replications were set up in swe- den in t e rst al o t e 1 30s (Lan let 1 34). n Poland,
ro enance lots sin a classic la o t were esta lis ed 30
ears later. owe er, t ese e eri ents tested onl art o t e seed ase (Barzda n 200 ). e oldest ro enance lots o sil er r were esta lis ed in o w in 1 61 ( nia 1 6
zeli ows i et al. 2011). e res lts o eas re ents and o ser ations cond cted nia (2006) and B (2007) s ow a reat de ree o aria ilit in t e anal sed o lation in terms of growth and qualitative characteristics.
e s ste o eld trials in ol in di erent en iron en- tal conditions to test populations representing a wide range of natural species allows us to gain a better understanding of the character of the genetic variation of adaptive traits than nder conditions o nat ral rowt (Ait en, annerz 2001).
according to the premises of the program testing the pro- en o orest trees, 37 ro en testin lots o 7 o la- tions and 2 ot er trees o sil er r ro all o r re ions of testing were established in Poland as of 2015, covering t e entire ran e o t is s ecies in t e co ntr ( i . 1). n test region iV (south-east 2), one population set is being tested
( ro en o 1 selected stands) in arallel wit t e national standard, regional standard and local standards (Fig. 2). the e eri ental la o t incl des o r trial lots a in di erent conditions o altit de, e os re and soil ertilit ( i . 3).
e st d s ects are assessed or t eir ada tation to ad er- se growing conditions, among other factors, which in the
resent e eri ent is e ressed a red ced s sce ti ilit to bud and needle damage due to freezing temperatures. the res lts resented are a on t e rst on t e assess ent o t e sensiti it o rs to late rosts. e testin ro ra oc sed on antitati e traits ( a or et al., 2004) in st d in t e ada-
tation o t e ro en o selected seed stands and ot er trees to di erent rowt conditions. or t is reason, t e eld trials assess onl s rin d rst, w ereas s sce ti ilit to rost da a e is assessed onl in t e e ent o its occ rrence.
Arti cial reezin tests co ined wit t eir eri cation in eld trials a e not een incl ded in t e testin ro ra , even for such a low temperature sensitive species as silver r ( olnic i 2003). e location o t e sil er r ro en test lots in so t eastern Poland ta es into acco nt t e ris o rost in o ntaino s areas eca se o t e con ration and co er o t e terrain (Ko ni s i 1 74). is was con r ed
t e eriodic occ rrence o late rosts at all test lots.
t dies on sil er r cond cted to date were ainl con- cerned wit learnin a o t t e siolo ical ec anis s o acquiring frost resistance and how this changes over time ( olnic i 2003 ar a 2004). e iss e o t e aria ilit in a sil er r o lation o its s sce ti ilit to da a e ro
late rosts is still ins cientl e lored ( an o ic et al.
2007). e ai o t is wor is to select o lations o sil er r t at can est ada t to t e rowin conditions o so t e- astern Poland and are c aracterised a si ni cantl i er resistance to late rosts in t eir rst ears o rowt .
2. Materials and methods
e tested ro en o selected r stands re resent o- pulations from eight districts of the regional Directorate of state Forests in Krosno (table. 1). the trial plots were esta- blished in 2009 in the districts of cisna (cis) – n49°09’60’’,
22 27 7 Ko a cza ( ) – 4 22 2 , 22 00 7 Bircza (B ) – 4 3 24 , 22 2 34 and Les o (L ) – n49°29’51’’, e22°17’36’’ (Fig. 2), each with an area of 225 m2 in a rando ised loc desi n wit o r re lications. All experimental plots were established under the cover of eu- ro ean larc lanted two ears earlier. a itat conditions at t e test lots are aried (res ecti el : – L św, – L św, B – Lw św L – Lw św), t eet t e tro ic requirements of the test species. in terms of regionalisation, the natural-forest areas are situated in the carpathian region, in t e esore ions o t e Prze s ie lands (B ), la ( ), t e Low Bieszczad (L ), t e i Bieszczad ( ) ( ielon , Klicz ows a 2012). eedlin s wit a co ered root s ste (1. / 1. K) were sed to esta lis t e cro and were planted in 1.5 × 1.5 m2. each population was represented
400 seedlin s (100 or eac re lication).
Table 1. characteristics of tested populations
Population iD Forest Base Material regional Directorate
of state Forest Forest District Provenance region
Bal2 11626 Krosno Bali r d 806
Bir1 27146 Krosno Bircza 804
Kra1 11270 Krosno Krasicz n 804
Kra2 11271 Krosno Krasicz n 804
les1 30320 Krosno Les o 806
les2 30326 Krosno Les o 806
les3 30327 Krosno Les o 806
les5 30329 Krosno Les o 806
lUt1 30205 Krosno L towis a 806
lUt2 30212 Krosno L towis a 806
lZD MP/2/45114/06 Kra w L Kr nica 803
nar 10995 Krosno narol 606
rYM1 36145 Krosno an w 806
rYM2 36146 Krosno an w 806
stU 10691 Krosno t osian 806
late frosts occurred twice (in 2009 and 2013) in the se en ears since t e tested cro s were lanted, ca sin da a e to t e a ical and lateral s oots o t e r. Varia ilit in the degree and nature of the damage was observed in terms of both the experimental plots as well as the tested o lations. e t e o da a e was deter ined i e- diatel a ter t e rost occ rrence. A t ree de ree scale was
sed to assess da a e d rin eld o ser ations to deter i- ne the extent of damage to the shoots (intact trees, dama- ed lateral s oots, da a ed a ical s oot). n t is st d , we anal sed t e o ser ations o da a e to t e lateral s oots and apical shoot.
in order to determine the proportion of trees expe- riencin di erent t es o da a e, we deter ined t e
re enc o s c da a e or indi id al ro enances. o deter ine t e ariation in t e t e o da a e a on t e test lots and tested o lations o r, we sed t e eneral linear odel, co inin a cross classi cation wit a ie- rarc ical one ( ta a 1 0): lot ear t e o da a e
o lation and t e e ect o nested loc s in t e interac-
tion o site ear: Figure 1. estin re ions or elected eed tands o sil er r:
i – Middle eastern, ii – south Western, iii – south eastern 1, iV – south eastern 2
Figure 2. location of tested population and experimental lots (B – Bircza orest istrict, L – Les o orest istrict, K – Ko a cza orest istrict, – isna Forest District)
Fijklm=µ + Si + Yj + FTk + Pl + SYij + SFTik +YFTjk+ SPil + YPjl + FTPkl + SYFTijk + SYPijl +SFTPikl + YFTPjkl + Bm(SYij) + Eijklm
where:
Fijkm – percentage of frost damage of site i, in ear j and t e of damage k,
µ – overall average, Si – effect of site i, Yj – e ect o ear j,
FTk – e ect o t e o da a e k, Pl – effect of population l,
SYij – interaction of site i and ear j,
SFTik – interaction of site i and t e o da a e k, SPil – interaction of site i and population l, YPjl – interaction o ear j and population l,
FTPkl – interaction o t e o da a e k and population l, YFTik – interaction o ear j and t e o da a e k,
SYFTijk – interaction of site i, ear j and t e o da a e k, SYPijl – interaction of site i, ear j and population l, SFTPikl – interaction of site i, t e o da a e k and popu- lation l,
YFTPjkl – interaction o ear j, t e o da a e k and popu- lation l
Bm(SYij) – e ect o nested loc m in the interaction, site i and ear j,
Eijklm – random error.
e anal sis was ased on data o tained ro t e Bliss transformation. homogeneous groups were determined
sin e s onestl i ni cant i erence ( ) test.
tatistical anal ses were er or ed sin t e L roce- dure of sas (sas institute 2011).
3. Results
e anal sis o ariance con r ed si ni cance di eren- ces in the rates of the occurrence of frost damage among the tested populations (p = 0.002), sites (p 0.001), t e ears 2009 and 2014 (p 0.001), and t e second and t ird t es o damage (p 0.001). At t e sa e ti e, t e si ni cance o t e
interaction was con r ed or: t e o da a e o lation, t e o da a e site, t e o da a e ear and site ear (in all cases p 0.001). e anal ses cond cted also con r-
ed t e si ni cance o t e interaction etween t e o rost da a e and ear o occ rrence and t e tested o lations wit t e site, res ecti el : p = 0.001 and p < 0.001 (table 2).
e anal sis o ariance de onstrated a si ni cant di - erence in t e re enc o t e incidence o t e second and t ird t es o da a e, w ic was con r ed in t e a orit o t e tested o lations. nl t e L o lation (t e na- tional standard) was c aracterised t e sa e n er o trees with damaged lateral shoots and apical shoot (Fig. 4).
For most of the remaining tested populations, the third da- a e t e (da a ed a ical s oot) was o ser ed si ni can- tl ore re entl t an t e second da a e t e (da a ed lateral s oots). e di erence in t e re enc o da a e t es etween t e L 1 and A o lations was not sta- tisticall si ni cant.
e anal sis o t e re enc o t e occ rrence o ot rost da a e t es o ser ed in 200 and 2014 s owed a different trend for damage to the apical shoots and lateral s oots. e re enc o t e second da a e t e (da a ed lateral s oots) was si ilar in ot late rost ears or ost of the tested populations. the proportion of damaged trees in 200 was si ni cantl reater t an in 2014 onl or t e lZD population (the national standard). at the same time, t e trees o t e K A1 o lation ad t e least re entl o ser ed second da a e t e ( i . ). e o osite trend was o nd or t e re enc o t e t ird da a e t e (da- a ed a ical s oot). All o lations s owed a si ni cantl reater re enc o da a e to t e a ical s oot ro t e late frost of 2009 compared to the 2014 frost. in the case of the 200 late rost, t e t ird da a e t e was least li e to a e been observed on the trees of the lZD population. the late rosts o 2014 ost ea il da a ed t e L 2 o lation, t t e di erence in t e re enc o da a e to t e re a- inin o lations (e ce t or L ) was not statisticall con- r ed. n 2014, t e re enc o da a e to t e a ical s oot o t e tested o lations was ore e enl distri ted and no si ni cant di erences were o nd a on o lations.
Figure 3. elevation gradient and geographical directions of exposure of experimental plots. MgZ – mountain grassland zone, DFZ – lower mountain decidous forest zone, FZ – foothills zone. other designation as in Figure 2.
4. Discussion
e ris o rost da a e de ends ot on t e increase in average annual temperature and the impact of climate c an e on t e incidence ( re enc and scale) o s ort ter te erat re ct ations (Be er et al., 1 ). Po lations of conifer species growing in variable mountain environ- ments exhibit greater interpopulation variation than popula-
tions rowin in a ore o o eneo s en iron ent (Ait en, hannerz 2001), so the selection of mountain populations of sil er r or resistance to rost da a e can ro a l i -
ro e t eir ada ti e otential ( olnic i, K c ci s i 2003).
Accordin a ora (1 ), t e enetic al e o sil er r from the Polish carpathians region provides the basis for attempting to select a population that is best able to adapt to rowin in o ntaino s areas c aracterised t e re ent Table 2. Anal sis o ariance
source of variation sum of squares Deegres of
freedom Mean square F-statistic p-value
e o rost da a e 6.58 1 6.58 367.09 <0.001
Population 0.62 14 0.04 2.47 0.002
site 3.29 3 1.10 61.26 <0.001
Year 36.32 1 36.32 2026.32 <0.001
e o rost da a e o lation 1.07 14 0.08 4.27 <0.001
e o rost da a e site 0.84 3 0.28 15.6 <0.001
e o rost da a e ear 34.46 1 34.46 1922.28 <0.001
Population × site 0.81 42 0.02 1.07 0.352
Po lation ear 0.32 14 0.02 1.27 0.218
ite ear 3.3104 3 1.10 61.55 <0.001
e o rost da a e o lation site 0.93 42 0.02 1.24 0.145
e o rost da a e o lation ear 0.80 14 0.06 3.17 0.001
e o rost da a e site ear 3.75 3 1.25 69.69 <0.001
Po lation site ear 0.31 42 0.01 0.42 0.999
Bloc (site ear) 1.46 24 0.06 3.39 <0.001
error 13.23 738 0.02
Figure 4. rans or ed a era e or re enc o late rost da a e o sil er r o lations ( ot ears) error ars eans e continuous line – 3rd da a e t e dotted line – 2nd da a e t e.
occurrence of late frosts. assessing the adaptive potential of this species to environmental conditions differing from its place of origin is based on observations conducted in pro-
enance areas and in e eri ental lots testin t e ro en o e cl ded seed stands and ot er trees (Barzda n 200 Kowal ows i 2013 rz szews a 2013). o ar, Poland has not conducted research on the population’s genetic va- ria ilit in ter s o rost ardiness nder controlled rowt conditions. Past st dies on t e ada ti e otential o sil er r populations from Poland examined characteristics providing indirect e idence o its s sce ti ilit to da a e, s c as e- i t, s r i a ilit and t e enolo o d de elo ent (Ke , a or 200 rz szews a 2010 zeli ows i et al.
2011). enetic aria ilit o rost ardiness as een directl con r ed or ot er coni er s ecies st dies ased ot on eld trials as well as arti cial reezin tests ( or lin 1 2 annerz 1 4 i son 1 4 Ait en, Ada s 1 7 Be - er et al. 1 olnic i, Kra 1 Andersson, edor o 2004; langvall 2011). the results of these studies suggest t at t e sil er r also as an ada ti e otential, conditioned eneticall , o resistance to tiss e da a e ca sed erio- dic frosts during the growing season. the temperature limit,
elow w ic trees are da a ed rost, de ends on an factors: the seasonal variation of the frost hardiness of trees, the duration of sustained frosts, sun exposure and the pace of temperature drops (hannerz 1994). Bearing in mind the
location of the experimental plots (Fig. 3), it should be assu- ed t at t ese actors stron l di erentiate t e i act o late rosts on t e t e and intensit o t e res ltin da a e.
e lac o onitorin o local weat er conditions does not allow t e ass tions listed a o e to e directl eri ed and t e se o lti ear eteorolo ical data in o ntaino s re- gions does not provide the basis for determining the actual intensit o t e rosts (Ait en, Ada s, 1 7). e aria ilit o t e intensit o late rosts related not onl to t e sites o t e e eri ental lots, t also to t e ear in w ic t e occ rred. idence o t is is seen in ot t e si ni cantl different numbers of damaged trees as well as the proportion of trees with damaged lateral shoots and apical shoot (Fig.
). A si ni cantl lower ro ortion o trees wit a da a ed apical shoot in 2014 compared to 2009 could have occurred eca se o ot a lower intensit o t e rost in 2014, as well as a lower s sce ti ilit o t e trees to da a e related to t e acclimatization of the tested population. the diverse respon- se o trees to rost co ld also a e een in enced t e ambient conditions occurring prior to the growing season, responsible for the timing of bud development (Dormling 1982). comparing the proportion of apical shoot damage a on t e tested o lations in ot anal sed ears allows t e statisticall si ni cant di erences etween o lations and etween t e ears o rost to e o ser ed. n 200 , t e lZD population, serving as the national standard in the pro-
Figure 5. rans or ed a era e or re enc of late frost damage (2nd da a e t e – side shoots, 3rd da a e t e – a ical s oots) o sil er r o lations in s se ent ears error ars eans e . ontin o s line means late frost in 2014; dotted line means late frost in 2009.
en testin ro ra , was t e ost resistant to t is t e o da a e. sce ti ilit to rost da a e at a enile a e is one o t e i ortant c aracteristics t at a a ect t e ia ilit o t e cro s. ed ced s sce ti ilit o L tree stand ro en to late rosts a indicate a i de-
ree o ada ti it in t is o lation. e inter o lation ariation o rost ardiness o ser ed in t e rst ear o rowt o t e trees was reatl red ced a ter si ears. t is true that the lZD population continued to be characte- rised a low de ree o a ical s oot da a e, t si ilar
al es were also reac ed o lations ro locations w ic were t e art est awa ro t e test sites: K A1 and A ( i . ). is attern is con r ed t e e- neral t eor c aracterizin local ro enances as s o ti- al o lations ( at as. eat an 1 2). Per a s t is observed trend was associated with a later seedling bud burst of the lZD population in 2009, while in 2014, the remaining populations acquired resistance to late frosts, reaching a height above which the phenomenon of frost no longer affects the development of apical buds. During research into the frost hardiness of spruce, Danusevicius and his team (1999) observed a greater degree of frost da a e in older trees. e e lained t is attern t e greater surface of the assimilation apparatus in older (hi- gher) plants leading to a loss of water.
e a t ors o t is st d elie e t at d e to t e a i- it o Anderson and edor o a s (2004) res lts w ic disc ss t e in ence o seedlin ei t on s sce ti ilit to rost da a e, t e st d s o ld ta e into acco nt ot an- titative traits as well as the adaptive potential of seedlings.
or t is reason, t e test res lts o t e ro en o selected sil er r tree stands and ot er trees s o ld also relate to both quantitative traits as well as their adaptive potential.
election or ada ti it ai s to de ne t e est trees in ter s of resistance to damage from frost and population, as well as to de ne t e tar et en iron ental conditions or t eir se (Ait en, annerz 2001). e resented a roac to selec- tin sil er r or o ntaino s conditions is sti ed d e to t e wide aria ilit o en iron ental conditions.
5. Conclusions
e o lation o sil er r in t e so t eastern testin (second) region most susceptible to damage from late frosts in t e rst ear o rowt was t e tree stand selected ro t e orest eri ental tation (L ) in Kr nica.
A ter si ears o rowt in t e so t eastern testin (second) re ion, t e ro en o tree stands selected ro t e orest istricts o Krasicz n and arol ( o. L P LP res ecti el 11270 and 10 ) e i ited t e lowest s sce - ti ilit to da a e ro late rosts.
Assessin t e co rse o a reaction to en iron ental sti- li o sil er r o lations nder o ntain cli ate con- ditions c aracterised re ent occ rrences o late rost requires the continuous monitoring of microclimate con- ditions occurring at the experimental plots.
Conflicts of interest
e a t ors declare t at no otential con icts o interest exist.
Acknowledgements and source of funding
e st d was nded t e tate orests ational o- rest oldin as art o researc ro ect BLP 37 .
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Authors’ contribution
.K. – data collection and its statistical anal sis, re ara- tion o t e an scri t, coordinatin t e wor z. . – initial preparation, data collection, text corrections; J.U. – statisti- cal anal sis o t e data, te t corrections P.P. – data collec- tion, text corrections; J.M. – concept and premises of the st d . . – carto ra ic editin , te t corrections.
ranslated : Bar ara Prz ls a
