Method notes, Exp II, “Genes vs. Species”

Jason Fridley and Phil Grime, Fall-Winter 2004 plus Fall 2005 update

 

See also details in Excel Design Spreadsheet.

 

Planting

 

Eight genotypes were selected for each of eight species, derived from planted material that had been continually propagated at Tapton Garden by Rosemary Booth (Booth and Grime 2003) since __.  Genotypes were originally collected by Booth and Grime from a 10 x 10 m area at Cressbrookdale.  Genotypes were subsequently subjected to molecular analysis by Whitlock (Whitlock in prep) and all 64 genotypes selected for the present study have been corroborated as unique genotypes by microsatellite analysis.  Genotypes had been propagated in a mix of compost and perlite and kept in both greenhouse and outdoor conditions, arranged in randomized blocks.

 

Genotypes were planted during a three-week period from mid-November to early-December 2004 by JDF and JPG, one genotype at a time and one species at a time.  All individuals of Festuca were planted first, followed by Koeleria, Helictotrichon, Carex caryophyllea, C. flacca, C. panicea, Campanula, and Succisa.  Individuals were prepared by splitting apart separate tussocks, potted genets, or spreading clumps (Succisa), and clipping above and belowground tissues to a standard biomass within species (typically 6-8 cm above- and belowground).  Roots were wetted prior to planting.  After planting, boxes were kept constantly moist and warm by heating cables (thermostat set to 10 C) and overhead plasting sheeting (briefly) to encourage root and shoot growth.

 

Sixty-four individuals were planted in each box in an equally spaced 8 x 8 grid.  For the 8 species x 8 genotype treatment, planting positions were determined randomly and each genotype was represented once in each box.  For the 8 spp/4 genotype and 4spp/8genotype treatments, each genotype was planted twice and replicate compositions were planted on each side of the box with respect to crevice location, but within a box side planting positions were random.  For the 6 other treatments, genotypes were represented by 4, 8, 16, 32, or 64 individuals and replicate compositions were planted in each quadrant of the box, but randomly within quadrants.

 

Soils

 

All soils were removed from the field, nearby the Buxton Climate Change Laboratory in Harpur Hill, Derbyshire.  Two locations were chosen to match the rendzina and podzol soil types occurring in or near the Cressbrookdale location.  The more calcicolous vegetation above the rendzina soil was chosen about halfway down the slope of a limestone dale at Buxton, while the podzol (acid) soil was chosen from an area near the top of an adjacent dale with more calcifuge vegetation (Vaccinium, Arrhenathrum?).  Rendzina was removed (turf vegetation down to bedrock, 20-30 cm deep), placed in plastic bags on site, hauled up with a wench system (acknowledgement: Andrew Askew!), and transported back to Tapton.  The soil was then dried a bit in greenhouses, mixed in a cement mixer and mixed further by hand to remove major rocks, root tussocks and large rhizomes, but was otherwise unamended (and was noticeably rich in earthworms).  Podzol was removed and treated in much the same way but the immediate surface turf was not bagged (soil depth 40-50 cm); few macrofauna were apparent.  Crevice soil (see below) was filled with a separately-removed rendzina, from the same location as the fresh rendzina but removed and stored as living turves at Tapton for __ years (8?).  Turves were allowed to dry out before soil was removed from roots with a pick, mixed in a wheelbarrow and placed down the PVC pipes by hand.

 

Crevices were assembled on boxes with soil already present.  To the fill the box with soil, first the bottom 5 cm over the whole box was filled with fresh rendzina.  Boxes were filled all at once with soil batches in aliquots.  On top of this 5 cm rendzina layer, boxes were further filled with a 5 cm deep checkerboard layout of fresh rendzina and podzol, separated with a cardboard divider (initially) to facilitate sharp soil boundaries.  The checkerboard layer divided each box into 4 quadrants and each soil type went into diagonal patches; for each box, the orientation of the patches with respect to the plot layout AND the crevice location was randomly determined.  Total soil depth (on non-crevice/rock side) was 10 cm initially and settled to 8-9 cm.

 

Box construction

 

Boxes (experimental unit) were constructed from tannalized timber with inner dimensions 55 x 55 x 12 cm.  Slats over the bottom left 2 cm wide gaps and were covered with water permeable nylon mesh to facilitate drainage.  Box soil structure was varied by allowing each box 4 soil “crevices” extending below the box, surrounded by 2 “rocks” that halved soil depth to 5 cm.  DIAGRAM.  Crevices were PVC pipes measuring 46 cm long x 3.2 cm in diameter, extending out from the bottom of the box and oriented horizontally in a line on one side of the box, __ from one end and equally spaced at 11 cm intervals.  On either side of the crevices were two pieces of tannalized lumber measuring 55 x 10 x 5 cm deep, inserted and nailed onto the box bottom, over the nylon mesh.  Such “rocks” were intended to provide further soil heterogeneity on the crevice portion of the box and increase drought stress to plants planted above them.  Soil structure was further complicated by a patchwork design of rendzina and podzol soil types (see above).

 

Plot layout

 

Fifty-four boxes were arranged along two rows of scaffolding containing 6 randomized blocks of 9 treatments.  Boxes were protected from herbivory and animal disturbance by mesh fencing.  [Treatments and box layout are detailed in excel sheet.]

 

Treatment Compositions

 

Species diversity treatments were chosen to be specific subsets of an eight species mixture: all 8 species; Fo, Km, Hp, Cc; and Festuca only.  Festuca ovina is one of the most widespread and abundant species in many types of British grassland including the limestone grassland at Cressbrookdale, is highly resistant to drought and grazing, and occurs on a wide range of soil types.  The other two species composition treatments represent increasing complexity from the Festuca monoculture: first two other very common tussock grasses and the clonal sedge Cc, followed by common subdominant components at Cressbrookdale, two forbs and two other sedges.

 

Genotypes of each of these species were chosen based on the 5-year results of their performance in the synthesized communities of Booth and Grime (2003).  Chosen genotypes represented a gradient of high to low abundance in 16-genotype treatments of the Booth and Grime experiment.  In the case of Cc and Cr, biotypes that were not verified as unique genotypes by Whitlock were excluded.  In the case of Succisa, genotypes were chosen largely on the basis of available stock material.

 

In contrast to species composition, genotype composition for each treatment was chosen randomly, with the following constraints:

 

Species

Genotypes / sp

How GD compositions assembled

1

1

Each of the 8 Festuca genotypes put in one box (here replication = 8)

4

1

Randomly select 6 of the 8 genotypes of each species, then randomly seed each into a replicate

8

1

Randomly select 6 of the 8 genotypes of each species, then randomly seed each into a replicate

1

4

Each of the 8 genotypes of Festuca represented 3 times over 6 reps, otherwise random

4

4

Each of the 8 genotypes of 4 species represented 3 times over 6 reps, otherwise random

8

4

Each of the 8 genotypes of 8 species represented 3 times over 6 reps, otherwise random

1

8

Exact replicates of 8-genotype mix (here only 4 replicates)

4

8

Exact replicates of 8-genotype mix

8

8

Exact replicates of 8-genotype mix

 

Maintenance

 

27 May 05      Box 11 four of the Cr3’s look desiccated and dying back.  Not replaced.  Box 9 disturbed by squirrels Cc2, Cc9 both replaced by material from greenhouse pots. Cr8 replaced by cutting both Booth and Grime box 18.

 

5 Oct 05         15 dead individuals replaced, including:

                        Box 3 Hp2

                        Box 6 Hp1

                        Box 8 Hp1

                        Box 7 Hp1

                        Box 9 Hp3

                        Box 13 Cc9 (2)

                        Box 18 Hp11

                        Box 20 Hp1 (2)

                        Box 22 Hp1

                        Box 34 Cp15

                        Box 38 Cp5

                        Box 28 Hp3

                        Box 45 Hp1

                        Campanula not replaced, due to difficulty determining whether actually “dead”.  Likely that most individuals will return in the spring.

 

First simulated grazing at 2.5 cm height applied on 13 July 2005, coinciding with Mark Bilton’s third census and Chris Bennett and Elizabeth Richell’s recording of reproductive and vegetative vigor with Fo.  Second clipping performed 28 Sept – Oct 4 2005.  All clippings saved for weighing by quadrant, and individuals of four species (Hp, Km, Cp, Cf) harvested by individual.  Other four species either have little biomass above cut (Sp, Cr, Cc) or have already been measured once (Fo).