Macroelement (N, P, K) Contents of Romulea columnae Seb. and Mauri Subsp. columnae During Vegetative and Generative Growth Phases
Tu— ba Kok,
Hamdi Guray Kutbay
Romulea columnae Seb. and Mauri subsp. columnae is a geophytic plant belonging to Iridaceae. In this study nitrogen (N), phosphorus (P) and potassium (K) analysis were carried out during vegetative and generative growth periods. It has been found that above ground parts of plant have higher macroelement concentrations as compared to below ground parts during vegetative growth period. However, below ground parts have higher macroelement concentrations during generative growth phase due merely to top senescence. In addition to this there were significant and mostly negative correlations between plant and soil macroelement concentrations.
Geophytic plants evade stress conditions such as shade, drought etc., by survival
in below ground organs and they exhibit, hereby, some features of sun plants
but also those of plants shaded habitats (Goryshina, 1972).
Geophytes are the plants in which the perennating bud is borne on a subterranean
storage organ and their annual growth cycle usually includes a dormant period.
The reserves in geophytic plants in their storage organ support leaf growth
at the beginning of the growing season and to a varying degree, also reproduction.
They also classified as spring ephemerals (Méndez,
1999; Lapointe, 2001). Spring ephemerals have a
very short epigeous (period during which shoots are present above ground) growth
period of 40-60 days in spring (Lapointe, 2001; Kilinc
et al., 2005). In addition to this they have also interesting phenological
properties such as flowering time ( spring or autumn), the presence of protantherous
and hysteranthous taxa etc., (Rawal et al., 1991;
Feller and Fischer, 1994).
Romulea columnae Seb. and Mauri subsp. columnae is a spring geophyte
belonging to Iridaceae and it is a Mediterranean element and it is a new record
for the study area (A6 square) according to the grid system of Davis
(1984). This species widely used as an ornamental plant. It consists of
some alkaloids in its bulbs most of them are not clearly identified (Baytop,
1984). So, that the bulbs of this taxa have continuously been exporting
to the abroad so that population density of this taxa have been decreased day
by day (Ekim et al., 2000).
In this study the redistribution of some macroelements ( N, P, K) between above
and below ground parts and the correlations between plant and soil during vegetative
and generative growth periods were examined to determine whether this species
followed the same pattern with the other spring ephemerals in terms of macronutrient
using strategy (i.e., top senescence phenomena) or not.
The concentrations of nitrogen, phosphorus and potassium seem to be more closely
controlled than other nutrients, which could reflect the specific amounts needed
for biochemical function (Canadell and Vilà, 1992).
MATERIALS AND METHODS
This study was carried out around Kurupelit region, near Bafra town which
is situated in the northern part of Turkey (36° 10 E, 41° 22
N) in 2001. Plant individuals were taken from open grassy places in a maquis
community dominated by Phillyrea latifolia L. at 150 m. The plants are
densely occured in this region. Outside this region plants were scattered due
to urbanization and overgrazing (Ekim et al., 2000).
Mean annual temperature and mean annual precipitation in the study area are
14.3°C and 712 mm, respectively. A xeric period which is one of the main
indicators of Mediterranean climate (Daget, 1977) was
observed from June to September in the study area. According to the Embergers
precipitation-temperature formula ([Q = 2000 x P/(M+m+546.4) (M-m)] where Q
is the precipitation-temperature index; M is maximum temperature and m is minimum
temperature, respectively) Q, M and m values are 129.3, 23.2 and 4.0° C,
respectively and the climate of the study area is humid Mediterranean (Daget,
1977; Ministry of Agriculture, 1994).
Ten plant individuals used for macroelement analysis in each of vegetative and generative growth phases. In other words, sampling was repeated twice during vegetative and generative growth phases. Phenological observations were also recorded.
Method of Chemical Analysis
Plant samples were harvested during vegetative and generative growth phases
and separated into above and below ground parts. After being washed in deionized
water the plant parts were dried at 70°C to the constant weight and grounded
in a Wiley mill and pass through a 20 mesh sieve. Nitrogen was determined by
the micro Kjeldahl method with a Kjeltec 1030 Analyser ( Tecator, Sweden) after
digesting the samples in concentrated H2SO4 with a selenium
catalyst. For P and K analysis plant specimens were wet ashed in concentrated
HNO3 and HClO4 and P was determined by using Jenway spectrophotometer
and K was determined by Petracourt PFP-7 flame photometer (Allen
et al., 1986).
Soil samples were collected during vegetative and generative growth phases
separately and soil and plant samples were taken simultaneously during vegetative
and generative growth phases. Soil samples were taken using a 7 cm diameter
auger to a depth of 30 cm. Ten soil cores were taken according to a fixed spatial
arrangement after the plant samples removed. Soil samples were air-dried and
sieved to pass through a 2 mm. mesh prior to analysis. Soil texture was determined
by Bouyoucous hydrometer method. pH values were measured in deionized water
(1:1). Total salinity (%) was determined by conductivity bridge apparatus. Soil
nitrogen (%) was determined by micro Kjeldahl method. Soil phosphorus (%) was
determined spectrophotometrically following the extraction by ammonium acetate.
Soil potassium (%) was determined by using a Petracourt PFP-7 flame photometer
after nitric acid wet digestion. Organic matter (%) and CaCO3 (%)
concentrations were determined by Walkley-Black method and Scheibler calcimeter
respectively (Bayrakli, 1987). The results of soil analysis
were explained according to Chapmann and Pratt (1973) and Bayrakli
The differences were assessed by one-way ANOVA test. Pearson correlation coefficients
were also calculated. Statistical analysis were performed using MINITAB software
package (Schaefer and Anderson, 1989).
Romulea columnae subsp. columnae is a protantherous taxa and
leaves appeared before the flowers. Leaves were appeared at the end of February.
The flowering time of R. columnae subsp. columnae is the second
half of March. In other words, R. columnae subsp. columnae sharply
switches from vegetative to reproductive growth. The appearance of fruits takes
place at the second half of April and at the first half of May the seeds are
The Results of Plant and Soil Analysis
Romulea columnae subsp. columnae occur on clay loamy soils.
Soil N and K concentrations were medium levels. However soil P concentrations
were low levels. Romulea columnae subsp. columnae grow on non-calcerous
and non-saline soils. Soil pH was slightly acidic during vegetative growth
phase, however neutral during generative growth phase. Organic matter levels
were high in both growth periods. (Table 1). Soil N, P and
K were decreased during generative growth phase. However, soil organic matter,
soil pH and CaCO3 were increased during generative growth phase (Table
Macroelement concentrations of above and below ground parts of R. columnae subsp. columnae during vegetative and generative growth phases were shown in Table 3 and 4. In vegetative growth period above ground parts have higher nutrient concentrations as compared to below ground parts However, below ground parts have higher nutrient concentrations during generative growth phase inversely (Table 3 and 4).
|| Mean values for soil parameters
|| The comparison between vegetative and generative growth periods
in respect to soil factors
|*: p< 05; **: p< 01; NS: Not Significant
|| Above ground macro element concentrations during vegetative
and generative growth phase
|| Below ground macroelement concentrations during vegetative
and generative growth phases
|| Statistical comparison of above and below ground macroelement
|*:p< 05; **: p< 01
There were significant differences between above and below ground macroelement
concentrations both vegetative and generative growth phases (Table
5). There were also significant and mostly negative correlations between
plant and soil macroelement concentrations in above and below ground parts in
both growth periods.
There were significant differences between two growth periods in respect to
soil N concentration, organic matter, pH and total salinity. The other soil
factors were not significantly changed (Table 2). Soil pH
and organic matter were inherently interrelated (Goldberg
1982). Increasing in pH during generative growth phase were related to the
return of dead organic matter consists of basic materials to the surface of
the soil (Singh, 1989).
Macroelement concentrations in above ground parts of R. columnae subsp.
columnae are higher than below ground parts during vegetative growth
phase (Table 3 and 4) due to the fast division
of meristematic cells in above ground parts. High nitrogen concentrations in
meristematic tissues were depend on the high protein content of that tissues
(Werger and Hirose, 1991; Moorby and
Beresford 1983). However, in generative growth period below ground parts
have higher nutrient concentrations as compared to above ground parts. Anderson
and Eickmeier (2000) reported that many species resorb nutrients from their
above ground parts back to below ground parts during senescence and the transfer
of nutrients is often associated with soil nutrient availability. During the
epigeous growth period, spring ephemerals accumulate mineral nutrients in their
below ground organs and develop the buds for the next years growth. Dormancy
is broken in autumn with bud and below ground growth that continues throughout
the winter at a very slow pace, due to low soil temperatures and this growth
period is called hypogeous growth, as it occurs below ground (Lapointe,
2001; Kilinc et al., 2005).
Similar results were also obtained in several other studies on geophytic plants
( Pirdal, 1989; Kutbay and Kilinç,
1993; Kutbay and Kilinç, 1995; Sahin
1998; Kutbay 1999). This situation is known as top
senescence (Leopold, 1980). In such plants the above ground
parts senesce completely and new shoots appear at the beginning of the next
season. The reserves in the vegetative storage organs allow a rapid growth during
initial phase ( Steinmann and Brandle, 1984; Nooden,
1984; Berchtold et al., 1993). Senescence
is an important process in the adaptation of higher plants to environmental
conditions. This is a well controlled process and it is not a passive decay
of a plant (Feller and Fischer, 1994). Senescence is
allowed to the optimum usage of macroelements for a plant (Jayasekera,
|| Pearson correlation coefficients between plant and soil macro
element concentrations in above ground parts
|** p< 01; NS: Not Significant
|| Pearson correlation coefficients between plant and soil macro
element concentrations in below ground parts
|* p< 05; ** p<01; NS:Not Significant
In addition to the top senescence monocotyledonous herbs have also adaptive
advantages as compared to dicotyledonous herbs. For example, above ground parts
of monocotyledonous herbs develop their leaves from a basal meristem. However,
dicotyledonous herbs develop their leaves from an apical meristem. As a result
of this meristematic tissues are at ground level in monocotyledonous herbs.
This means that the benefit of a basal meristem at ground level, in terms of
effective using of macroelements especially nitrogen, rapid transfer of nutrients
between above and below ground parts, providing protection against damage through
grazing, fire etc., (Werger and Hirose, 1991).
Canadell and Vilà (1992) found significant and
negative correlation coefficients between plant and soil nutrients. Knops
and Koenig (1997) found positive significant correlations between soil nitrogen
and phosphorus and foliar nitrogen and phosphorus. Powers
(1984) and Johnson et al. (1987) also found
positive correlation coefficients between soil and plant nutrient levels. Mostly
negative correlation coefficients were obtained between plant and soil macroelement
concentrations in above and below ground parts of R. columnae subsp.
columnae (Table 6 and 7).
These results suggest that soil nitrogen, phosphorus and potassium may influence
plant nutrient levels in most plants. However, there was species-specific differences
in this respect. Ecosystems dominated by short or long-lived species develop
soil over a multi-generational influencing soil to a small but eventually important
degree (Knops and Koenig 1997). Top senescence is an
important strategy to the adaptation of geophytic plants to environmental conditions
and main aim of this strategy is effective using of nutrients. The results of
the present study can be evaluated in cultivation of R.columnae subsp.
columnae. Future research should focus on top senescence in geophytic
plants to a more precise explanation the nutrient patterns during vegetative
and generative growth phases.
Allen, S.E., N.M. Grimshaw, J.A. Parkinson, C. Quarmby and J.D. Roberts, 1986.
Chemical Analysis. In: Methods in Plant Ecology, Chapman, S.B. (Ed.). Blackwell Scientific Publications, Oxford, pp: 411-466
Anderson, W. and W.G. Eickmeier, 2000.
Nutrient resorption in Claytonia virginica
L.: Implications for deciduous forest nutrient cycling. Can. J. Bot., 78: 832-839.Direct Link |
Bayrakli, F., 1987.
Plant and soil analysis. University of Ondokuz Mayis, Faculty of Agriculture Publications, Samsun, pp: 199.
Baytop, T., 1984.
Phytotherapy in Turkey From Past to Future. Department of Pharmacy Publications, Istanbul, pp: 520
Berchtold, A., J.M. Besson and U. Feller, 1993.
Effects of fertilization levels in two farming systems on senescence and nutrient contents in potato leaves. Plant Soil, 154: 81-88.Direct Link |
Canadell, J. and M. Vila, 1992.
Variation in tissue element concentration in Quercus ilex
L. over a range of different soils. Vegetatio, 99: 273-282.Direct Link |
Davis, P.H., 1984.
Flora of Turkey and the East Aegean Islands. Vol. 8, Edinburgh University Press, Edinburgh
Daget, P.H., 1977.
Mediterranean bioclimate: Analysing of climate types according to the Emberger system. Vegetatio, 34: 87-103.
Ekim, T., M. Koyuncu, M. Vural, H. Duman, Z. Aytac and N. Adiguzel, 2000.
Red Data Book of Turkish Plants (Pteridophyta
). 1st Edn., TDKA and Van Centennial University Press, Ankara
Feller, U. and A. Fischer, 1994.
Nitrogen metabolism in senescing leaves. Crit. Rev. Plant Sci., 13: 241-273.CrossRef |
Goldberg, D.E., 1982.
The distribution of evergreen and deciduous trees relative to soil type: An example from the Sierra Madre, Mexico and a general model. Ecology, 63: 942-951.
Goryshina, T.K., 1972.
Ecophysiological investigations on ephemeroid plants in forest-steppe zone of Central Russia. Oecol. Plant., 7: 241-258.
Jayasekera, R., 1993.
Interelement relationships in leaves of tropical montane trees. Vegetatio, 109: 145-151.Direct Link |
Johnson, J.E., C.L. Haag, J.G. Bockheim and G.G. Erdmann, 1987.
Soil-site relationships and soil characteristics associated with even-aged red maple (Acer rubrum
) stands in Wisconsin and Michigan. For. Ecol. Man., 21: 75-89.
Knops, J.M.H. and W.D. Koenig, 1997.
Soil fertility and leaf nutrients of sympatric evergreen and deciduous species of Quercus
in central coastal California. Plant Ecol., 130: 121-131.Direct Link |
Kutbay, H.G. and M. Kilinc, 1993.
An autecological study on Leucojum aestivum
L. Turk. J. Bot., 17: 1-4.
Kutbay, H.G. and M. Kilinc, 1995.
An autecological study on Galanthus rizehensis
Stern (Amaryllidaceae) Turk. J. Bot., 19: 235-240.
Kutbay, H.G., 1999.
Top senescence in Sternbergia lutea
(L.) Ker-Gawl ex Sprengel and Narcissus tazetta
L. subsp. tazetta
. Turk. J. Bot., 23: 127-131.Direct Link |
Lapointe, L., 2001.
How phenology influences physiology in deciduous forest spring ephemerals. Physiol. Plant., 113: 151-157.Direct Link |
Leopold, A.C., 1980.
Ageing and Senescence in Plant Development. In: Senescence in Plants, Thimann, K.V. (Ed.). CRC Press, California
Mendez, M., 1999.
Effects of sexual reproduction on growth and vegetative propagation in the perennial geophyte Arum italicum
(Araceae). Plant Biol., 1: 115-120.Direct Link |
Ministry of Agriculture, 1994.
Mean and Extreme Temperature and Rainfall Values. General Directorate of Meteorology, Ankara, Turkey
Moorby, J. and R.T. Beresford, 1983.
Mineral Nutrition and Growth. In: Encylopedia of Plant Physiology, Lauchli, A. and R.L. Bieleski (Eds.). Springer, Berlin, pp: 481-527
Nooden, L.D., 1984.
Integration of soybean pod development and monocarpic senescence. Physiol. Plant., 62: 273-284.
Pirdal, M., 1989.
An autecological study on Asphodelus aestivus
Brot. Turk. J. Bot., 13: 89-101.
Powers, R.F., 1984.
Estimating Soil Nitrogen Availability Through Soil and Foliar Analysis. In: Forest Soils and Treatment Impacts, Stone, E.L. (Ed.). Knoxville, Tennessee, pp: 353-379
Rawal, R.S., N.S. Bankoti, S.S. Samant and Y.P.S. Pangtey, 1991.
Phenology of tree layer species from the timberline around Kumaun in central Himalaya, India. Vegetatio, 93: 108-118.Direct Link |
Sahin, N.F., 1998.
Morphological, anatomical and physiological studies on Galanthus ikariae
Baker and Galanthus rizehensis
Stern (Amaryllidaceae) grown around NE Turkey. Pak. J. Bot., 30: 117-131.Direct Link |
Schaefer, R.L. and R.B. Anderson, 1989.
The Student Edition of MINITAB. User' s Manual Addison Wesley Publishing Company Inc., New York
Singh, L., 1989.
Dry matter and nutrient inputs through litter fall in a dry tropical forest of India. Vegetatio, 98: 129-140.
Steinmann, F. and R. Brandle, 1984.
Carbohydrate and protein metabolism in the rhizomes of bulrush (Schoenoplectus lacustris
(L.) Palla) in relation to natural development of the whole plant. Aquat. Bot., 19: 53-63.
Werger, M.J.A. and T. Hirose, 1991.
Leaf nitrogen distribution and whole canopy photosynthetic carbon gain in herbaceous stands. Vegetatio, 97: 11-20.Direct Link |
Kilinc, M., A. Bilgin, E. Yalcin and H.G. Kutbay, 2005.
Macroelement (N, P, K) contents of Arum euxinum
R. mill during vegetative and generative growth phases. Pak. J. Biol. Sci., 8: 267-272.CrossRef | Direct Link |