| Literature DB >> 24864293 |
N Leksrisompong1, H Romero-Sanchez2, E O Oviedo-Rondón1, J Brake3.
Abstract
A study was conducted to examine how 2 feeder space allocations during the rearing period followed by 2 feeder space allocations after photostimulation and 2 female feeding to peak programs (fast or slow) affected female broiler breeder reproductive performance and mortality. Sixteen pens of 76 breeder females each were equipped with either 4 tube feeders with a 132 cm circumference pan (7.0 cm/female) or 6 feeders (10.4 cm/female) to 21 wk of age. Thereafter, 64 females were moved to breeding pens, photostimulated, and fed sex-separate from either 3 (6.2 cm/female) or 5 (10.3 cm/female) feeders with either fast or slow feeding to peak feeding programs applied to complete a 2 × 2 × 2 factorial design. Seven males that were separately reared in a similar manner were added per pen. Individual female BW was determined at 6, 20, and 32 wk of age and BW uniformity assessed. Greater feeder space during rearing increased BW at 32 wk of age, whereas greater feeder space during lay or slow feeding to peak decreased BW at 32 wk. There were no differences in BW uniformity. Hens from the 10.4 to 10.3 cm/female combination produced a significantly greater number of eggs as compared with the 7.0 to 10.3 cm/female and 10.4 to 6.2 cm/female combinations with the 7.0 to 6.2 cm/female combination intermediate. Percentage hen-day egg production of the 10.4 to 10.3 cm/female combination hens was significantly greater than all other combinations. Livability was improved in the 10.4 to 10.3 cm/female combination relative to the 7.0 to 10.3 cm/female combination with the others intermediate. The fast feeding to peak program increased yolk weight as well as yolk:albumen ratio at 28 and 30 wk of age, but egg weight did not differ. These data indicated that increased or decreased feeder space between the growing and laying periods did not affect broiler breeder female BW, uniformity, egg weight, fertility, or hatchability. The 10.3 cm/female laying feeder space exhibited the best hen-day egg production in combination with 10.4 cm/pullet rearing but not with 7.0 cm/pullet rearing space. In a similar manner, hen mortality was greater in the 7.0 to 10.3 cm/female feeder space combination that the 10.4 to 10.3 cm/female combination.Entities:
Keywords: body weight; broiler breeder; egg production; feeder space; uniformity
Mesh:
Year: 2014 PMID: 24864293 PMCID: PMC4988534 DOI: 10.3382/ps.2013-03277
Source DB: PubMed Journal: Poult Sci ISSN: 0032-5791 Impact factor: 3.352
Figure 1.Ross 708SF pullets received feed according to this feeding program from placement to 21 wk of age. The figure shows the daily feeding amounts from which the 4/3 restricted feeding amounts were calculated. Pullets were fed from 3 wk of age on Monday, Wednesday, Friday, and Saturday each week until 21 wk of age when they were moved to the laying house.
Composition of starter (1 to 6 wk), grower (7 to 25 wk), and breeder (26 to 64 wk) diets
| Item | Starter diet | Grower diet | Breeder diet |
|---|---|---|---|
| Ingredient (%) | |||
| Corn | 64.30 | 65.49 | 66.73 |
| Soybean meal (48% CP) | 22.17 | 16.53 | 19.17 |
| Wheat bran | 7.98 | 12.61 | 3.00 |
| Dicalcium phosphate | 2.04 | 1.99 | 1.70 |
| Limestone | 1.13 | 1.02 | 5.94 |
| Mineral premix1 | 0.20 | 0.20 | 0.20 |
| Vitamin premix2 | 0.10 | 0.10 | 0.10 |
| Salt | 0.63 | 0.64 | 0.50 |
| Coccidiostat (Amprol) | 0.05 | 0.05 | 0.05 |
| | 0.07 | 0.04 | 0.06 |
| Selenium premix | 0.10 | 0.10 | 0.10 |
| Lysine HCl | — | 0.01 | — |
| Choline chloride | 0.20 | 0.20 | 0.20 |
| Poultry fat | 1.00 | 1.00 | 2.23 |
| Antibiotic | 0.03 | 0.03 | 0.03 |
| Totals | 100.00 | 100.00 | 100.00 |
| Calculated analysis3 | |||
| CP (%) | 17.00 | 15.00 | 15.00 |
| ME (kcal/g) | 2.93 | 2.93 | 2.93 |
| Lysine (%) | 0.88 | 0.75 | 0.77 |
| Methionine + cysteine (%) | 0.71 | 0.63 | 0.63 |
| Calcium (%) | 0.95 | 0.90 | 2.70 |
| Available phosphorus (%) | 0.45 | 0.45 | 0.40 |
1Mineral premix contained the following per kilogram of diet: manganese, 120 mg; zinc, 120 mg; iron, 80 mg; copper, 10 mg; iodine, 2.5 mg; and cobalt, 1.0 mg.
2Vitamin premix contained the following per kilogram of diet: vitamin A, 13,200 IU; cholecalciferol, 4,000 IU; vitamin E, 66 IU; vitamin B12, 34.6 µg; riboflavin, 13.2 mg; niacin, 110 mg; pantothenic acid, 22 mg; vitamin K, 4 mg; folic acid, 2.2 mg; thiamine, 4 mg; pyridoxine, 8 mg; and biotin, 252 µg.
3Data presented on an as-is basis.
Figure 2.Ross 708SF hens received restricted feed amounts daily according to these 2 feeding to peak programs. Either a fast feed to peak program from 21 to 28 wk of age or a slow feed to peak program from 21 to 29 wk of age was employed. All the hens received the same feed rate previously (Figure 1) and thereafter as shown above. Squares represent the fast feeding to peak program, and circles represent the slow feeding to peak program.
Broiler breeder female BW as affected by broiler breeder female feeder space allocation during growing and laying periods, and feed increase rate from photostimulation to peak egg production, as well as interactions among these factors
| Feeder space/female | BW (kg) | ||||
|---|---|---|---|---|---|
| Growing2 (cm) | Laying3 (cm) | Peak feed increase1 | 6 wk of age | 20 wk of age | 32 wk of age |
| 7.0 | 0.789 | 2.263 | 3.463 | ||
| 10.4 | 0.800 | 2.263 | 3.507 | ||
| 0.525 | 0.981 | 0.072 | |||
| 6.2 | 0.791 | 2.277 | 3.512a | ||
| 10.3 | 0.798 | 2.250 | 3.458b | ||
| 0.685 | 0.128 | 0.033 | |||
| Fast | 0.794 | 2.258 | 3.539A | ||
| Slow | 0.796 | 2.269 | 3.432B | ||
| SEM | 0.0354 | 0.0244 | 0.0215 | ||
| 0.920 | 0.505 | 0.009 | |||
| 7.0 | 6.2 | 0.803 | 2.279 | 3.483 | |
| 7.0 | 10.3 | 0.776 | 2.247 | 3.444 | |
| 10.4 | 6.2 | 0.793 | 2.274 | 3.542 | |
| 10.4 | 10.3 | 0.807 | 2.253 | 3.472 | |
| 0.231 | 0.754 | 0.486 | |||
| 7.0 | Fast | 0.786 | 2.255 | 3.531 | |
| 7.0 | Slow | 0.794 | 2.271 | 3.395 | |
| 10.4 | Fast | 0.803 | 2.261 | 3.546 | |
| 10.4 | Slow | 0.798 | 2.266 | 3.468 | |
| 0.690 | 0.725 | 0.200 | |||
| 6.2 | Fast | 0.795 | 2.272 | 3.575 | |
| 6.2 | Slow | 0.802 | 2.281 | 3.449 | |
| 10.3 | Fast | 0.794 | 2.244 | 3.502 | |
| 10.3 | Slow | 0.790 | 2.256 | 3.414 | |
| SEM | 0.0466 | 0.0316 | 0.0297 | ||
| 0.749 | 0.942 | 0.397 | |||
a,bMeans in a column that possess different superscripts differ significantly (P ≤ 0.05).
A,BMeans in a column that possess different superscripts differ significantly (P ≤ 0.01).
1Feed increase rates (see Figure 2) from photostimulation to peak egg production.
2Feeder space allocations of 7.0 or 10.4 cm per each of 76 pullets during growing.
3Feeder space allocations of 6.2 or 10.3 cm per each of 64 hens during laying.
4SEM for n = 8 pens with 76 pullets weighed individually per pen.
5SEM for n = 8 pens with 64 hens weighed individually per pen.
6SEM for n = 4 pens with 76 pullets weighed individually per pen.
7SEM for n = 4 pens with 64 hens weighed individually per pen.
Coefficient of variation of broiler breeder female BW as affected by broiler breeder female feeder space allocation during growing and laying periods, and feed increase rate from photostimulation to peak egg production, as well as interactions among these factors
| Feeder space/female | CV (%) | ||||
|---|---|---|---|---|---|
| Growing2 (cm) | Laying3 (cm) | Peak feed increase1 | 6 wk of age | 20 wk of age | 32 wk of age |
| 7.0 | 13.40 | 13.39 | 9.57 | ||
| 10.4 | 14.36 | 14.57 | 9.41 | ||
| | 0.249 | 0.292 | 0.853 | ||
| 6.2 | 13.80 | 14.24 | 9.60 | ||
| 10.3 | 13.95 | 13.72 | 9.39 | ||
| | 0.878 | 0.684 | 0.805 | ||
| Fast | 13.58 | 13.98 | 9.31 | ||
| Slow | 14.17 | 13.98 | 9.68 | ||
| SEM | 0.584 | 0.744 | 0.595 | ||
| | 0.449 | 0.999 | 0.665 | ||
| 7.0 | 6.2 | 13.02 | 13.73 | 9.81 | |
| 7.0 | 10.3 | 13.77 | 13.06 | 9.33 | |
| 10.4 | 6.2 | 14.58 | 14.75 | 9.39 | |
| 10.4 | 10.3 | 14.13 | 14.39 | 9.44 | |
| 0.559 | 0.885 | 0.755 | |||
| 7.0 | Fast | 13.10 | 13.65 | 9.20 | |
| 7.0 | Slow | 13.69 | 13.14 | 9.95 | |
| 10.4 | Fast | 14.06 | 14.31 | 9.41 | |
| 10.4 | Slow | 14.65 | 14.82 | 9.42 | |
| | 0.962 | 0.638 | 0.666 | ||
| 6.2 | Fast | 13.39 | 14.53 | 9.19 | |
| 6.2 | Slow | 14.21 | 13.95 | 10.01 | |
| 10.3 | Fast | 13.77 | 13.43 | 9.42 | |
| 10.3 | Slow | 14.13 | 14.01 | 9.35 | |
| SEM | 0.826 | 1.046 | 0.837 | ||
| | 0.824 | 0.595 | 0.608 | ||
1Feed increase rates (see Figure 2) from photostimulation to peak egg production.
2Feeder space allocations of 7.0 or 10.4 cm per each of 76 pullets during growing.
3Feeder space allocations of 6.2 or 10.3 cm per each of 64 hens during laying.
4SEM for n = 8 pens with 76 pullets per pen weighed and CV calculated on a pen basis.
5SEM for n = 8 pens with 64 pullets per pen weighed and CV calculated on a pen basis.
6SEM for n = 4 pens with 76 pullets per pen weighed and CV calculated on a pen basis.
7SEM for n = 4 pens with 64 pullets per pen weighed and CV calculated on a pen basis.
Egg weight (EW), percentage egg components, and yolk:albumen ratio at 28 wk of age as affected by broiler breeder female feeder space allocation during growing and laying periods, and feed increase rate from photostimulation to peak egg production, as well as interactions among these factors
| Feeder space/female | |||||||
|---|---|---|---|---|---|---|---|
| Growing2 (cm) | Laying3 (cm) | Peak feed increase1 | Egg weight (EW; g) | Yolk weight (% of EW) | Egg shell (% of EW) | Albumen (% of EW) | Yolk:albumen ratio (g:g) |
| 7.0 | 51.79 | 26.75 | 9.48 | 63.77 | 0.42 | ||
| 10.4 | 51.71 | 26.73 | 9.52 | 63.74 | 0.42 | ||
| 0.805 | 0.930 | 0.275 | 0.884 | 0.99 | |||
| 6.2 | 51.90 | 26.79 | 9.40B | 63.80 | 0.42 | ||
| 10.3 | 51.60 | 26.69 | 9.61A | 63.71 | 0.42 | ||
| 0.350 | 0.559 | 0.001 | 0.565 | 0.811 | |||
| Fast | 51.80 | 26.99a | 9.51 | 63.49b | 0.43a | ||
| Slow | 51.70 | 26.49b | 9.50 | 64.02a | 0.41b | ||
| | 0.745 | 0.019 | 0.642 | 0.013 | 0.017 | ||
| SEM4 | 0.22 | 0.12 | 0.02 | 0.12 | 0.01 | ||
| 7.0 | 6.2 | 51.62 | 26.98 | 9.40 | 63.62ab | 0.43 | |
| 7.0 | 10.3 | 51.96 | 26.52 | 9.56 | 63.92ab | 0.42 | |
| 10.4 | 6.2 | 52.19 | 26.61 | 9.40 | 63.99a | 0.42 | |
| 10.4 | 10.3 | 51.23 | 26.85 | 9.65 | 63.50b | 0.43 | |
| 0.067 | 0.078 | 0.216 | 0.044 | 0.074 | |||
| 7.0 | Fast | 52.04 | 26.91 | 9.53ab | 63.55 | 0.43 | |
| 7.0 | Slow | 51.54 | 26.58 | 9.43b | 63.98 | 0.42 | |
| 10.4 | Fast | 51.57 | 27.07 | 9.49b | 63.44 | 0.43 | |
| 10.4 | Slow | 51.86 | 26.39 | 9.56a | 64.05 | 0.41 | |
| 0.237 | 0.340 | 0.041 | 0.598 | 0.436 | |||
| 6.2 | Fast | 52.17 | 27.02 | 9.39 | 63.59 | 0.43 | |
| 6.2 | Slow | 51.63 | 26.56 | 9.42 | 64.02 | 0.42 | |
| 10.3 | Fast | 51.43 | 26.96 | 9.64 | 63.40 | 0.43 | |
| 10.3 | Slow | 51.77 | 26.41 | 9.58 | 64.01 | 0.41 | |
| | 0.193 | 0.802 | 0.234 | 0.603 | 0.764 | ||
| SEM5 | 0.31 | 0.17 | 0.03 | 0.17 | 0.02 | ||
a,bMeans in a column that possess different superscripts differ significantly (P ≤ 0.05).
A,BMeans in a column that possess different superscripts differ significantly (P ≤ 0.01).
1Feed increase rates (see Figure 2) from photostimulation to peak egg production.
2Feeder space allocations of 7.0 or 10.4 cm per each of 76 pullets during growing.
3Feeder space allocations of 6.2 or 10.3 cm per each of 64 hens during laying.
4SEM for n = 8 pens with mean of approximately 30 eggs weighed per pen.
5SEM for n = 4 pens with mean of approximately 30 eggs weighed per pen.
Egg weight (EW) percentage egg components and yolk:albumen ratio at 30 wk of age as affected by broiler breeder female feeder space allocation during growing and laying periods, and feed increase rate from photostimulation to peak egg production, as well as interactions among these factors
| Feeder space/female | |||||||
|---|---|---|---|---|---|---|---|
| Growing2 (cm) | Laying3 (cm) | Peak feed increase1 | EW (g) | Yolk weight (% of EW) | Egg shell (% of EW) | Albumen (% of EW) | Yolk:albumen ratio (g:g) |
| 7.0 | 54.84 | 28.03 | 9.24 | 62.73 | 0.45 | ||
| 10.4 | 54.98 | 27.86 | 9.16 | 62.98 | 0.44 | ||
| 0.761 | 0.350 | 0.127 | 0.202 | 0.38 | |||
| 6.2 | 55.01 | 27.91 | 9.19 | 62.90 | 0.45 | ||
| 10.3 | 54.80 | 27.97 | 9.22 | 62.81 | 0.45 | ||
| 0.638 | 0.714 | 0.585 | 0.624 | 0.787 | |||
| Fast | 55.10 | 28.24A | 9.16 | 62.60b | 0.45A | ||
| Slow | 54.72 | 27.64B | 9.25 | 63.11a | 0.44B | ||
| SEM4 | 0.30 | 0.12 | 0.03 | 0.12 | 0.01 | ||
| 0.404 | 0.007 | 0.084 | 0.020 | 0.005 | |||
| 7.0 | 6.2 | 54.60 | 28.23a | 9.25 | 62.52b | 0.45a | |
| 7.0 | 10.3 | 55.08 | 27.82ab | 9.23 | 62.94ab | 0.44ab | |
| 10.4 | 6.2 | 55.43 | 27.60b | 9.13 | 63.28a | 0.44b | |
| 10.4 | 10.3 | 54.52 | 28.13ab | 9.20 | 62.68b | 0.45ab | |
| 0.143 | 0.022 | 0.394 | 0.020 | 0.022 | |||
| 7.0 | Fast | 55.13 | 28.36 | 9.17 | 62.47 | 0.46 | |
| 7.0 | Slow | 54.55 | 27.69 | 9.31 | 63.00 | 0.44 | |
| 10.4 | Fast | 55.06 | 28.13 | 9.14 | 62.73 | 0.45 | |
| 10.4 | Slow | 54.89 | 27.59 | 9.19 | 63.22 | 0.44 | |
| 0.638 | 0.716 | 0.368 | 0.920 | 0.928 | |||
| 6.2 | Fast | 55.35 | 28.20 | 9.19 | 62.61 | 0.45 | |
| 6.2 | Slow | 54.68 | 27.63 | 9.18 | 63.19 | 0.44 | |
| 10.3 | Fast | 54.85 | 28.29 | 9.12 | 62.59 | 0.45 | |
| 10.3 | Slow | 54.76 | 27.66 | 9.31 | 63.03 | 0.44 | |
| SEM5 | 0.43 | 0.17 | 0.05 | 0.18 | 0.02 | ||
| | 0.522 | 0.716 | 0.064 | 0.687 | 0.703 | ||
a,bMeans in a column that possess different superscripts differ significantly (P ≤ 0.05).
A,BMeans in a column that possess different superscripts differ significantly (P ≤ 0.01).
1Feed increase rates (see Figure 2) from photostimulation to peak egg production.
2Feeder space allocations 7.0 or 10.4 cm per each of 76 pullets during growing.
3Feeder space allocations 6.2 or 10.3 cm per each of 64 hens during laying.
4SEM for n = 8 pens with mean of approximately 40 eggs weighed per pen.
5SEM for n = 4 pens with mean of approximately 40 eggs weighed per pen.
Broiler breeder egg production and female mortality from 24 to 64 wk as affected by broiler breeder female feeder space allocation during growing and laying periods, and feed increase rate from photostimulation to peak egg production, as well as interactions among these factors
| Feeder space/female | Egg production | ||||
|---|---|---|---|---|---|
| Growing2 (cm) | Laying3 (cm) | Peak feed increase1 | Eggs per hen housed (n) | Hen-day production (%) | Female mortality (%) |
| 7.0 | 172 | 62.3 | 8.8 | ||
| 10.4 | 175 | 63.2 | 7.8 | ||
| 0.326 | 0.077 | 0.525 | |||
| 6.2 | 172 | 62.2b | 8.2 | ||
| 10.3 | 175 | 63.3a | 8.4 | ||
| 0.195 | 0.037 | 0.525 | |||
| Fast | 173 | 62.7 | 9.0 | ||
| Slow | 174 | 62.8 | 7.6 | ||
| SEM4 | 2.4 | 0.7 | 1.3 | ||
| 0.922 | 0.741 | 0.382 | |||
| 7.0 | 6.2 | 175AB | 62.9B | 6.6ab | |
| 7.0 | 10.3 | 169B | 61.7B | 10.9a | |
| 10.4 | 6.2 | 168B | 61.5B | 9.8ab | |
| 10.4 | 10.3 | 181A | 65.0A | 5.9b | |
| 0.004 | 0.001 | 0.024 | |||
| 7.0 | Fast | 171 | 62.2 | 10.9 | |
| 7.0 | Slow | 173 | 62.4 | 6.6 | |
| 10.4 | Fast | 176 | 63.1 | 7.0 | |
| 10.4 | Slow | 174 | 63.3 | 8.6 | |
| 0.440 | 0.951 | 0.083 | |||
| 6.2 | Fast | 172 | 62.6 | 9.8 | |
| 6.2 | Slow | 171 | 61.8 | 6.6 | |
| 10.3 | Fast | 174 | 62.8 | 8.2 | |
| 10.3 | Slow | 176 | 63.9 | 8.6 | |
| SEM5 | 3.4 | 0.9 | 1.8 | ||
| | 0.583 | 0.070 | 0.270 | ||
a,bMeans in columns that possess different superscripts differ significantly (P ≤ 0.05).
A,BMeans in columns that possess different superscripts differ significantly (P ≤ 0.01).
1Feed increase rates (see Figure 2) from photostimulation to peak egg production.
2Feeder space allocations of 7.0 or 10.4 cm per each of 76 pullets during growing.
3Feeder space allocations of 6.2 or 10.3 cm per each of 64 hens during laying.
4SEM for n = 8 pens of 64 hens per pen.
5SEM for n = 4 pens of 64 hens per pen.
Broiler breeder fertility, hatchability, and embryonic mortality from 26 to 63 wk of age from weekly set of 60 eggs as affected by broiler breeder female feeder space allocation during growing and laying periods, and feed increase rate from photostimulation to peak egg production, as well as interactions among these factors
| Feeder space/female | Incubation variable2 (%) | |||||||
|---|---|---|---|---|---|---|---|---|
| Hatchability of | ||||||||
| Growing3 (cm) | Laying4 (cm) | Peak feed increase1 | Fertility | Fertile eggs | Total eggs | Early dead5 | Late dead6 | |
| 7.0 | 95.79 | 94.74 | 90.75 | 3.40 | 1.98 | |||
| 10.4 | 95.38 | 95.05 | 90.67 | 3.15 | 1.69 | |||
| 0.621 | 0.212 | 0.919 | 0.181 | 0.408 | ||||
| 6.2 | 95.17 | 94.86 | 90.29 | 3.43 | 1.72 | |||
| 10.3 | 96.00 | 94.92 | 91.13 | 3.12 | 1.89 | |||
| 0.334 | 0.684 | 0.383 | 0.120 | 0.712 | ||||
| Fast | 95.46 | 94.87 | 90.63 | 3.30 | 1.83 | |||
| Slow | 95.71 | 94.91 | 90.80 | 3.25 | 1.84 | |||
| | 0.873 | 0.976 | 0.880 | 0.802 | 0.973 | |||
| 7.0 | 6.2 | 95.73 | 94.87 | 90.83 | 2.65 | 1.88 | ||
| 7.0 | 10.3 | 95.85 | 94.60 | 90.68 | 3.15 | 1.89 | ||
| 10.4 | 6.2 | 94.61 | 94.85 | 89.75 | 2.77 | 1.56 | ||
| 10.4 | 10.3 | 96.16 | 95.24 | 91.59 | 2.48 | 1.88 | ||
| 0.364 | 0.455 | 0.322 | 0.071 | 0.815 | ||||
| 7.0 | Fast | 95.59 | 94.87 | 90.69 | 2.91 | 1.80 | ||
| 7.0 | Slow | 95.99 | 94.61 | 90.81 | 2.88 | 1.96 | ||
| 10.4 | Fast | 95.34 | 94.98 | 90.56 | 2.46 | 1.62 | ||
| 10.4 | Slow | 95.43 | 95.12 | 90.78 | 2.79 | 1.82 | ||
| 0.658 | 0.589 | 0.956 | 0.253 | 0.987 | ||||
| 6.2 | Fast | 94.89 | 94.73 | 89.90 | 2.58 | 1.80 | ||
| 6.2 | Slow | 95.46 | 95.00 | 90.68 | 2.84 | 1.64 | ||
| 10.3 | Fast | 96.04 | 95.12 | 91.35 | 2.79 | 1.63 | ||
| 10.3 | Slow | 95.96 | 94.73 | 90.91 | 2.83 | 2.15 | ||
| 0.904 | 0.338 | 0.604 | 0.575 | 0.373 | ||||
1Feed increase rates (see Figure 2) from photostimulation to peak egg production.
2Categorical analysis does not generate SEM.
3Feeder space allocations of 7.0 or 10.4 cm per each of 76 pullets during growing.
4Feeder space allocations of 6.2 or 10.3 cm per each of 64 hens during laying.
5Embryos that died from 1 to 7 d.
6Embryos that died after 7 d.
Figure 3.Percentage female mortality during the laying period as affected by female feeding space. Circles represent a small decrease in feeder space (7.0–6.2 cm/female); squares represent an increase in feeder space (7.0–10.3 cm/female); triangles represent a decrease in feeder space (10.4–6.2 cm/female); and diamonds represent a small decrease in feeder space (10.4–10.3 cm/female) at photostimulation.